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<mods:title id="EC6505E59448D09F812D4ABA67920DEE">Atlas of the skull bones of Tyrannosaurus rex</mods:title>
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<mods:namePart id="0CA3B98E335A4334366D6334081CABB2">Peter Larson</mods:namePart>
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<mods:namePart id="3A05D53CE64783FDA682A1BDD2DC5C97">Peter Larson</mods:namePart>
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<mods:namePart id="63C3FD396D9A97A15DEC303525A89320">Kenneth Carpenter</mods:namePart>
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<mods:title id="CE3FB5C7168A9722BAFBA52F9D376E29">Tyrannosaurus rex, the tyrant king</mods:title>
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<taxonomicName id="4C0B60E3E372D45AFF688BC7FE121689" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[165,466,709,745]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
Tyrannosaurus 
<emphasis id="B97FC772E372D45AFE698BC7FE121689" box="[420,466,709,745]" italics="true" pageId="0" pageNumber="233">rex</emphasis>
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was described by Osborn in a series of papers at the beginning of the last century (1905, 1906, 1912, 1916).
</paragraph>
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<subSubSection id="C31148EBE372D45AFBCC8BFAFADA1370" pageId="0" pageNumber="233" type="discussion">
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His work was primarily based on 3 specimens: the holotype (
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), the holotype for Dynamosaurus (
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, which was synonymized with 
<taxonomicName id="4C0B60E3E372D45AFE2E8A9BFDFC17DD" authorityName="Osborn" authorityYear="1905" box="[483,572,921,957]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
<emphasis id="B97FC772E372D45AFE2E8A9BFDFC17DD" box="[483,572,921,957]" italics="true" pageId="0" pageNumber="233">T. rex</emphasis>
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in 1906, and 
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, which provided much of the description of the skull. Interestingly, the designation of 
<materialsCitation id="3B63113DE372D45AFB3B8ACFFF351049" ID-GBIF-Occurrence="3396384333" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761" pageId="0" pageNumber="233" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
has recently come under scrutiny; it may actually represent a second, unnamed species of 
<taxonomicName id="4C0B60E3E372D45AFE3A8D3BFD26103D" authorityName="Osborn" authorityYear="1905" box="[503,742,1081,1117]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E372D45AFE3A8D3BFD26103D" box="[503,742,1081,1117]" italics="true" pageId="0" pageNumber="233">Tyrannosaurus</emphasis>
</taxonomicName>
(P. Larson this volume). 
<bibRefCitation id="EF9A6691E372D45AFBB08D3AFAB8103B" author="Molnar, R. E." box="[1149,1400,1079,1116]" journalOrPublisher="Palaeontographica Abteilung A" pageId="0" pageNumber="233" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar’s (1991)</bibRefCitation>
treatise on the skull added greatly to Osborn’s earlier descriptive work, and it also contained a section on arthrology. Molnar studied 
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, 
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, 
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, and 
<materialsCitation id="3B63113DE372D45AFD668DDBFC50109C" ID-GBIF-Occurrence="3396384366" box="[683,912,1240,1277]" collectionCode="SDSM" pageId="0" pageNumber="233" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
. In 1998 and in this volume, Molnar increased our understanding of skull mechanics and musculature. This was followed by 
<bibRefCitation id="EF9A6691E372D45AFDD68C46FCDF1108" author="Brochu, C. A." box="[539,799,1348,1384]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="0" pageNumber="233" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu’s (2003)</bibRefCitation>
description of 
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, including computed tomography of the articulated skull, presented significant new data and interpretations. Several other important contributions to the skull of tyrannosaurids include 
<bibRefCitation id="EF9A6691E372D45AFCD98CE3FC0F1265" author="Carr, T. D." box="[788,975,1505,1541]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="0" pageNumber="233" pagination="497 - 520" part="19" refId="ref5496" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19 (3): 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr (1999)</bibRefCitation>
on craniofacial ontogeny, 
<bibRefCitation id="EF9A6691E372D45AFF658F1AFE49125C" author="Currie, P. J." box="[168,393,1560,1596]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="0" pageNumber="233" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie (2003)</bibRefCitation>
on cranial anatomy of tyrannosaurid dinosaurs (particularly 
<taxonomicName id="4C0B60E3E372D45AFF6A8F4EFEB81210" authorityName="Cope" authorityYear="1866" box="[167,376,1612,1648]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E372D45AFF6A8F4EFEB81210" box="[167,376,1612,1648]" italics="true" pageId="0" pageNumber="233">Gorgosaurus</emphasis>
</taxonomicName>
and 
<emphasis id="B97FC772E372D45AFE1C8F4EFD241210" box="[465,740,1612,1648]" italics="true" pageId="0" pageNumber="233">
<taxonomicName id="4C0B60E3E372D45AFE1C8F4EFD121210" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[465,722,1612,1648]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
),
</emphasis>
<bibRefCitation id="EF9A6691E372D45AFD3F8F4EFBF61210" author="Currie, P. J. &amp; Hurum, J. H. &amp; Sabath, K." box="[754,1078,1612,1648]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="0" pageNumber="233" pagination="227 - 234" part="48" refId="ref5704" refString="Currie, P. J., Hurum, J. H., and Sabath, K. 2003. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontologia Polonica 48 (2): 227 - 234" title="Skull structure and evolution in tyrannosaurid dinosaurs" type="journal article" year="2003">Currie et al. (2003)</bibRefCitation>
on skull structure, 
<bibRefCitation id="EF9A6691E372D45AFF6A8F80FDA412C6" author="Hurum, J. H. &amp; Sabath, K." box="[167,612,1666,1702]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="0" pageNumber="233" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath (2003)</bibRefCitation>
on a comparison of 
<taxonomicName id="4C0B60E3E372D45AFC0A8F83FB5112C5" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" box="[967,1169,1665,1701]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E372D45AFC0A8F83FB5112C5" box="[967,1169,1665,1701]" italics="true" pageId="0" pageNumber="233">Tarbosaurus</emphasis>
</taxonomicName>
and 
<emphasis id="B97FC772E372D45AFB248F82FED912BB" italics="true" pageId="0" pageNumber="233">
<taxonomicName id="4C0B60E3E372D45AFB248F82FED312BB" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
,
</emphasis>
<bibRefCitation id="EF9A6691E372D45AFEE48FB5FCC912BB" author="Carr, T. D. &amp; Williamson, T. E." box="[297,777,1719,1755]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="233" pagination="479 - 523" part="142" refId="ref5527" refString="Carr, T. D., and Williamson, T. E. 2004. Diversity of Late Maastrichtian Tryrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 142: 479 - 523." title="Diversity of Late Maastrichtian Tryrannosauridae (Dinosauria: Theropoda) from western North America" type="journal article" year="2004">Carr and Williamson (2004)</bibRefCitation>
on tyrannosaurid diversity, and 
<bibRefCitation id="EF9A6691E372D45AFAE28FB7FEAC1371" author="Carr, T. D. &amp; Williamson, T. E. &amp; Schwimrner, D. R." journalOrPublisher="Journal of Vertebrate Paleontology" pageId="0" pageNumber="233" pagination="119 - 143" part="25" refId="ref5570" refString="Carr, T. D., Williamson, T. E., and Schwimrner, D. R. 2005. A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama. Journal of Vertebrate Paleontology 25 (1): 119 - 143." title="A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama" type="journal article" year="2005">Carr et al. (2005)</bibRefCitation>
on a description of a new tyrannosaurid from Alabama.
</paragraph>
</subSubSection>
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<paragraph id="8BB41B60E372D45AFF3D8E20FB5B1D44" blockId="0.[164,1408,703,2505]" pageId="0" pageNumber="233">
Although several new 
<taxonomicName id="4C0B60E3E372D45AFDBD8E20FC601325" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[624,928,1825,1862]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
<emphasis id="B97FC772E372D45AFDBD8E20FC601325" box="[624,928,1825,1862]" italics="true" pageId="0" pageNumber="233">Tyrannosaurus rex</emphasis>
</taxonomicName>
skulls are now available for study, by far the most significant is the disarticulated, undistorted skull of 
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(see N. L. Larson this volume). For the first time, the individuals bones of the skull can be described and illustrated for 
<taxonomicName id="4C0B60E3E372D45AFB8E8EC3FAB31384" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1091,1395,1984,2021]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
<emphasis id="B97FC772E372D45AFB8E8EC3FAF71385" box="[1091,1335,1985,2021]" italics="true" pageId="0" pageNumber="233">Tyrannosaurus</emphasis>
rex
</taxonomicName>
. This specimen was discussed by 
<bibRefCitation id="EF9A6691E372D45AFD198EF4FB501C7A" author="Hurum, J. H. &amp; Sabath, K." box="[724,1168,2038,2074]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="0" pageNumber="233" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath (2003)</bibRefCitation>
in their comparison of 
<taxonomicName id="4C0B60E3E372D45AFE928129FD511C2F" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[351,657,2091,2127]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
<emphasis id="B97FC772E372D45AFE928129FD511C2F" box="[351,657,2091,2127]" italics="true" pageId="0" pageNumber="233">Tyrannosaurus rex</emphasis>
</taxonomicName>
and 
<emphasis id="B97FC772E372D45AFD208128FBF71C2E" box="[749,1079,2090,2126]" italics="true" pageId="0" pageNumber="233">
<taxonomicName id="4C0B60E3E372D45AFD208128FBF21C2E" baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[749,1074,2090,2126]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
.
</emphasis>
<bibRefCitation id="EF9A6691E372D45AFB818128FAEA1C2E" author="Smith, J. B." box="[1100,1322,2090,2126]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="0" pageNumber="233" pagination="865 - 887" part="25" refId="ref6264" refString="Smith, J. B. 2005. Heterodonty in Tyrannosaurus rex: implications for the taxonomic and systematic utility of theropod dentitions. Journal of Vertebrate Paleontology 25 (4): 865 - 887" title="Heterodonty in Tyrannosaurus rex: implications for the taxonomic and systematic utility of theropod dentitions" type="journal article" year="2005">Smith (2005)</bibRefCitation>
also utilized 
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in his work describing the heterodont dentition of 
<taxonomicName id="4C0B60E3E372D45AFAD48162FAB31CE3" authorityName="Osborn" authorityYear="1905" box="[1305,1395,2143,2180]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
T. 
<emphasis id="B97FC772E372D45AFA8E815DFAB31CE3" box="[1347,1395,2143,2179]" italics="true" pageId="0" pageNumber="233">rex</emphasis>
</taxonomicName>
<emphasis id="B97FC772E372D45AFABE815DFAB71CE3" box="[1395,1399,2143,2179]" italics="true" pageId="0" pageNumber="233">.</emphasis>
This subject was also discussed in some detail in Osborn (1916), 
<bibRefCitation id="EF9A6691E372D45AFB328196FECD1C8F" author="Molnar, R. E." journalOrPublisher="Palaeontographica Abteilung A" pageId="0" pageNumber="233" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
, and 
<bibRefCitation id="EF9A6691E372D45AFEA181C9FD9E1C8F" author="Brochu, C. A." box="[364,606,2251,2287]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="0" pageNumber="233" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003)</bibRefCitation>
. P. Larson (this volume) has utilized this specimen in his discussion of an avianlike kinetic palate in 
<taxonomicName id="4C0B60E3E372D45AFBF68002FB571D44" authorityName="Osborn" authorityYear="1905" box="[1083,1175,2304,2340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
.
</paragraph>
<paragraph id="8BB41B60E372D45AFF3E8035FAD91DA4" blockId="0.[164,1408,703,2505]" pageId="0" pageNumber="233">
Here, I present the first in-depth supplemental description of the skull bones and illustrate them in multiple views in the supplemental CD-ROM, based primarily on 
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, with observations of other specimens.
</paragraph>
<paragraph id="8BB41B60E372D45AFA06833CF88F1E10" blockId="0.[1483,1871,2622,2672]" box="[1483,1871,2622,2672]" pageId="0" pageNumber="233">
Dermal Skull 
<emphasis id="B97FC772E372D45AF917833CF88F1E10" box="[1754,1871,2622,2672]" italics="true" pageId="0" pageNumber="233">Bones</emphasis>
</paragraph>
<paragraph id="8BB41B60E372D45BFF6A8345FC301694" blockId="0.[166,1405,2625,2722]" lastBlockId="1.[665,1913,183,2670]" lastPageId="1" lastPageNumber="234" pageId="0" pageNumber="233">
Premaxillae: The premaxillae of 
<materialsCitation id="3B63113DE372D45AFD008344FCA71E0A" ID-GBIF-Occurrence="3396384345" box="[717,871,2630,2666]" collectionCode="BHI" pageId="0" pageNumber="233" specimenCode="BHI 3033">BHI 3033</materialsCitation>
are typical for 
<taxonomicName id="4C0B60E3E372D45AFBAC8344FABB1E0A" authorityName="Osborn" authorityYear="1906" box="[1121,1403,2630,2666]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
and for 
<taxonomicName id="4C0B60E3E372D45AFEE4837EFD9A1EC0" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[297,602,2684,2720]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="233" phylum="Chordata" rank="species" species="rex">
Tyrannosaurus 
<emphasis id="B97FC772E372D45AFDE4837EFD9A1EC0" box="[553,602,2684,2720]" italics="true" pageId="0" pageNumber="233">rex</emphasis>
</taxonomicName>
<emphasis id="B97FC772E372D45AFD97837EFDA01EC0" box="[602,608,2684,2720]" italics="true" pageId="0" pageNumber="233">,</emphasis>
bearing 4 alveoli. Details not discussed by Osborn (1905, 1906, 1912), 
<bibRefCitation id="EF9A6691E373D45BFB8989BEFAF71480" author="Molnar, R. E." box="[1092,1335,188,224]" journalOrPublisher="Palaeontographica Abteilung A" pageId="1" pageNumber="234" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
, 
<bibRefCitation id="EF9A6691E373D45BFA8389BEF9821480" author="Brochu, C. A." box="[1358,1602,188,224]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="1" pageNumber="234" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003)</bibRefCitation>
, and 
<bibRefCitation id="EF9A6691E373D45BF96B89BEFC421574" author="Hurum, J. H. &amp; Sabath, K." journalOrPublisher="Acta Palaeontologia Polonica" pageId="1" pageNumber="234" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath (2003)</bibRefCitation>
, but visible in 
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, include a scalloped region of ridges on the articular surface shared by the right and left premaxillae. This scalloped region is centrally located dorsoventrally and is at the anterior border of the suture. Although nearly 80% of this suture is smooth, ligaments attached to the scalloped area could have restricted lateral movement along this surface, in effect bonding the 2 premaxillae into a single structural unit. Below this scalloped area, the premaxillae separate, leaving a cleft in the articulation. The significance of this cleft is unclear, and it is not seen in the articulated 
<materialsCitation id="3B63113DE373D45BFC0B8B9BFB0F16DD" ID-GBIF-Occurrence="3396384362" box="[966,1231,665,701]" collectionCode="FMNH" pageId="1" pageNumber="234" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
(
<bibRefCitation id="EF9A6691E373D45BFB278B9BFA0516DD" author="Brochu, C. A." box="[1258,1477,665,701]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="1" pageNumber="234" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu 2003</bibRefCitation>
), a robust morphotype (P. Larson this volume).
</paragraph>
<paragraph id="8BB41B60E373D45BFD2B8A06F93F17F3" blockId="1.[665,1913,183,2670]" pageId="1" pageNumber="234">The right premaxilla shows erosion surrounding the anterior portion of the fourth alveolus. This erosion is associated with the deposition of spongy bone. Osteomyelitis is the probable cause of this pathology.</paragraph>
<paragraph id="8BB41B60E373D45BFD2B8AA6FACD10FD" blockId="1.[665,1913,183,2670]" pageId="1" pageNumber="234">The palatal surfaces of the premaxillae also bear a feature analogous to the interdental plates Osborn 1912) of the maxillae and dentaries. These interdental plates are found at the junctions of the alveoli. They are depressed from the palatal surface of the premaxillae and separated from one another at the base by large nutrient foramina.</paragraph>
<paragraph id="8BB41B60E373D45BFD2A8DAEFB231226" blockId="1.[665,1913,183,2670]" pageId="1" pageNumber="234">
The articular surface of the premaxillary-maxillary suture is smooth, interrupted (in the center of the lateral aspect) only by the opening for the subnarialis foramen (
<bibRefCitation id="EF9A6691E373D45BFBC98C1AFB71115C" author="Carr, T. D." box="[1028,1201,1304,1340]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="234" pagination="497 - 520" part="19" refId="ref5496" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19 (3): 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr 1999</bibRefCitation>
; 
<bibRefCitation id="EF9A6691E373D45BFB0B8C1AFA61115C" author="Brochu, C. A." box="[1222,1441,1304,1340]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="1" pageNumber="234" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu 2003</bibRefCitation>
), whose border is shared by the 2 elements. The premaxillary-nasal articulation is also quite smooth, allowing dorsoventral movement while restricting anteroposterior movement. It is probable that the premaxillae unit was at least passively kinetic, movable relative to the maxillae and nasals, providing shock-absorbing benefits for the premaxillary teeth.
</paragraph>
<paragraph id="8BB41B60E373D45BFD278F5EFAC513E5" blockId="1.[665,1913,183,2670]" pageId="1" pageNumber="234">
Maxillae: The maxillae of 
<materialsCitation id="3B63113DE373D45BFB058F55FAA2121B" ID-GBIF-Occurrence="3396384307" box="[1224,1378,1623,1659]" collectionCode="BHI" pageId="1" pageNumber="234" specimenCode="BHI 3033">BHI 3033</materialsCitation>
bear 11 alveoli. This number is shared with 
<materialsCitation id="3B63113DE373D45BFCA68F8EFBD612D0" ID-GBIF-Occurrence="3396384327" box="[875,1046,1676,1712]" collectionCode="MOR" pageId="1" pageNumber="234" specimenCode="MOR 980">MOR 980</materialsCitation>
, although to date, all other 
<taxonomicName id="4C0B60E3E373D45BFA2E8F8EF8CB12D0" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1507,1803,1676,1712]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
specimens have 12 (P. Larson this volume). The 2 maxillae of 
<materialsCitation id="3B63113DE373D45BF9FB8FC0F90D1286" ID-GBIF-Occurrence="3396384365" box="[1590,1741,1730,1766]" collectionCode="BHI" pageId="1" pageNumber="234" specimenCode="BHI 3033">BHI 3033</materialsCitation>
articulate near the front of the palate in a series of overlapping ridges not mentioned by 
<bibRefCitation id="EF9A6691E373D45BFD528E29FC43132F" author="Molnar, R. E." box="[671,899,1835,1871]" journalOrPublisher="Palaeontographica Abteilung A" pageId="1" pageNumber="234" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
or 
<bibRefCitation id="EF9A6691E373D45BFC758E29FB62132F" author="Brochu, C. A." box="[952,1186,1835,1871]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="1" pageNumber="234" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003)</bibRefCitation>
. This is similar to the condition in 
<taxonomicName id="4C0B60E3E373D45BF91B8E29FC8413E5" authority="(Hurum and Sabath 2003)" baseAuthorityName="Maleev" baseAuthorityYear="1955" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B97FC772E373D45BF91B8E29FC8413E5" italics="true" pageId="1" pageNumber="234">Tarbosaurus bataar</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9A6691E373D45BFC978E63FB3613E5" author="Hurum, J. H. &amp; Sabath, K." box="[858,1270,1889,1925]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="1" pageNumber="234" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath 2003</bibRefCitation>
).
</paragraph>
<paragraph id="8BB41B60E373D45BFD268E95F9BA1C3B" blockId="1.[665,1913,183,2670]" pageId="1" pageNumber="234">
In 
<emphasis id="B97FC772E373D45BFCDA8E94FBD213DA" box="[791,1042,1942,1978]" italics="true" pageId="1" pageNumber="234">
<taxonomicName id="4C0B60E3E373D45BFCDA8E94FBCC13DA" authorityName="Osborn" authorityYear="1905" box="[791,1036,1942,1978]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
,
</emphasis>
the antorbital fossa is a very deep depression in even the smallest individuals (i.e., 
<materialsCitation id="3B63113DE373D45BFB6F8EC9FA87138F" ID-GBIF-Occurrence="3396384342" box="[1186,1351,1995,2031]" collectionCode="BHI" pageId="1" pageNumber="234" specimenCode="BHI 4100">BHI 4100</materialsCitation>
and 
<materialsCitation id="3B63113DE373D45BFA648EC9F947138F" ID-GBIF-Occurrence="3396384301" box="[1449,1671,1995,2031]" collectionCode="LACM" pageId="1" pageNumber="234" specimenCode="LACM 23845">LACM 2345</materialsCitation>
). The ventral border of the antorbital fossa and the antorbital fenestra are coincident along much of the ventral border of the antorbital fenestra.
</paragraph>
<paragraph id="8BB41B60E373D458FD268169FAB31576" blockId="1.[665,1913,183,2670]" lastBlockId="2.[171,1415,183,2728]" lastPageId="2" lastPageNumber="235" pageId="1" pageNumber="234">
In most tyrannosaurs, like 
<taxonomicName id="4C0B60E3E373D45BFB608169FABB1CEF" authorityName="Cope" authorityYear="1866" box="[1197,1403,2155,2191]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E373D45BFB608169FABB1CEF" box="[1197,1403,2155,2191]" italics="true" pageId="1" pageNumber="234">Gorgosaurus</emphasis>
</taxonomicName>
(TCM 2001.89.1; 
<bibRefCitation id="EF9A6691E373D45BF97A8169F8A31CEF" author="Carr, T. D." box="[1719,1891,2155,2191]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="234" pagination="497 - 520" part="19" refId="ref5496" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19 (3): 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr 1999</bibRefCitation>
), 
<taxonomicName id="4C0B60E3E373D45BFD5281A0FC5A1CA6" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[671,922,2210,2246]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E373D45BFD5281A0FC5A1CA6" box="[671,922,2210,2246]" italics="true" pageId="1" pageNumber="234">Daspletosaurus</emphasis>
</taxonomicName>
(RTMP 91.36.500; 
<bibRefCitation id="EF9A6691E373D45BFB2881A0FA701CA6" author="Currie, P. J." box="[1253,1456,2210,2246]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="1" pageNumber="234" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
), 
<emphasis id="B97FC772E373D45BFA0681A0F9331CA6" box="[1483,1779,2210,2246]" italics="true" pageId="1" pageNumber="234">Appalachiosaurus</emphasis>
(RMM 6670; 
<bibRefCitation id="EF9A6691E373D45BFCC881D7FBCD1C99" author="Carr, T. D. &amp; Williamson, T. E. &amp; Schwimrner, D. R." box="[773,1037,2261,2297]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="234" pagination="119 - 143" part="25" refId="ref5570" refString="Carr, T. D., Williamson, T. E., and Schwimrner, D. R. 2005. A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama. Journal of Vertebrate Paleontology 25 (1): 119 - 143." title="A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama" type="journal article" year="2005">Carr et al. 2005</bibRefCitation>
), and 
<taxonomicName id="4C0B60E3E373D45BFBB881D7FA9E1C99" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[1141,1374,2261,2297]" pageId="1" pageNumber="234">Nanotyrannus</taxonomicName>
(CMNH 7541, BMRP 2001.4.1; 
<bibRefCitation id="EF9A6691E373D45BFD6D8009FC071D4F" author="Bakker, R. T. &amp; Williams, M. &amp; Currie, P." box="[672,967,2315,2351]" journalOrPublisher="Hunteria" pageId="1" pageNumber="234" pagination="26" part="1" refId="ref5414" refString="Bakker, R. T., Williams, M., and Currie, P. 1988. Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana. Hunteria 1 (5): 26." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana" type="journal article" year="1988">Bakker et al. 1988</bibRefCitation>
; Larson in press), the antorbital fossa is shallower than 
<taxonomicName id="4C0B60E3E373D45BF8958009FD101D05" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="species" species="rex">
<emphasis id="B97FC772E373D45BF8958009F8B01D4F" box="[1880,1904,2315,2351]" italics="true" pageId="1" pageNumber="234">T.</emphasis>
rex
</taxonomicName>
, and a thin ridge of bone (the ventral antorbital maxillary ridge) rises along the dorsal margin of the posteroventral extension of the maxilla. It extends well past the last alveolus and passes under the jugal where it articulates to the maxilla. In 
<emphasis id="B97FC772E373D45BFBDE80DDF8EE1E63" box="[1043,1838,2527,2563]" italics="true" pageId="1" pageNumber="234">
<taxonomicName id="4C0B60E3E373D45BFBDE80DDFB241E63" authorityName="Cope" authorityYear="1866" box="[1043,1252,2527,2563]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
, 
<taxonomicName id="4C0B60E3E373D45BFB3880DDFA301E63" authority=", Appalachiosaurus, and Nanotyrannus" authorityName="Appalachiosaurus, and Nanotyrannus" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[1269,1520,2527,2563]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
, 
<taxonomicName id="4C0B60E3E373D45BFA3280DDF8E81E63" box="[1535,1832,2527,2563]" pageId="1" pageNumber="234">Appalachiosaurus</taxonomicName>
,
</emphasis>
and 
<taxonomicName id="4C0B60E3E373D45BFD528316FC491E58" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[671,905,2580,2616]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="234" phylum="Chordata" rank="genus">Nanotyrannus</taxonomicName>
, the ventral border of the antorbital fenestra and the antorbital fossa are not coincident. The constriction at the base of the ventral antorbital maxillary ridge forms the ventral border of the antorbital fossa, and the top of the ridge is the ventral border of the antorbital fenestra (Larson in press)
</paragraph>
<paragraph id="8BB41B60E370D458FF3B882AFA8B16DF" blockId="2.[171,1415,183,2728]" pageId="2" pageNumber="235">
The maxillary fenestra (the second antorbital fenestra of Osborn 1912) in 
<taxonomicName id="4C0B60E3E370D458FF13885FFE0D15E1" authorityName="Osborn" authorityYear="1905" box="[222,461,349,385]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
<materialsCitation id="3B63113DE370D458FE11885EFDBE15E0" ID-GBIF-Occurrence="3396384351" box="[476,638,348,384]" collectionCode="BHI" pageId="2" pageNumber="235" specimenCode="BHI 3033">BHI 3033</materialsCitation>
) and 
<taxonomicName id="4C0B60E3E370D458FD25885EFC6D15E0" authority="(Hurum and Sabath 2003)" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" box="[744,941,348,384]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E370D458FD25885EFC6D15E0" box="[744,941,348,384]" italics="true" pageId="2" pageNumber="235">Tarbosaurus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9A6691E370D458FC06885EFAB615E0" author="Hurum, J. H. &amp; Sabath, K." box="[971,1398,348,384]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="2" pageNumber="235" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath 2003</bibRefCitation>
) contacts the anterior and posterior border of the antorbital fossa. An additional opening between the maxillary and antorbital fenestrae is found on both maxillae of 
<materialsCitation id="3B63113DE370D458FE0188F9FDAE167F" ID-GBIF-Occurrence="3396384303" box="[460,622,507,543]" collectionCode="BHI" pageId="2" pageNumber="235" specimenCode="BHI 3033">BHI 3033</materialsCitation>
, but is not seen in other specimens of 
<taxonomicName id="4C0B60E3E370D458FB3488F9FA94167E" authorityName="Osborn" authorityYear="1905" box="[1273,1364,506,543]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="species" species="rex">
T. 
<emphasis id="B97FC772E370D458FAE988F8FA94167E" box="[1316,1364,506,542]" italics="true" pageId="2" pageNumber="235">rex</emphasis>
</taxonomicName>
<emphasis id="B97FC772E370D458FA9988F8FA98167E" box="[1364,1368,506,542]" italics="true" pageId="2" pageNumber="235">.</emphasis>
A similar opening is seen in a 
<taxonomicName id="4C0B60E3E370D458FD518B32FCA21634" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" box="[668,866,560,596]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E370D458FD518B32FCA21634" box="[668,866,560,596]" italics="true" pageId="2" pageNumber="235">Tarbosaurus</emphasis>
</taxonomicName>
specimen, 
<emphasis id="B97FC772E370D458FBE18B32FB501634" box="[1068,1168,560,596]" italics="true" pageId="2" pageNumber="235">ZPAL</emphasis>
MgD-I/4, described by 
<bibRefCitation id="EF9A6691E370D458FE978B65FCC816EA" author="Hurum, J. H. &amp; Sabath, K." box="[346,776,614,651]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="2" pageNumber="235" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath (2003)</bibRefCitation>
. This opening may be the result of predepositional weathering and breakage of an extremely thin area of bone.
</paragraph>
<paragraph id="8BB41B60E370D458FF388BD3FC471054" blockId="2.[171,1415,183,2728]" pageId="2" pageNumber="235">A long, narrow, and deep abrasion is incised into the lateral aspect of the left maxilla. This gouge is oriented dorsoventrally and is directly above the third maxillary tooth near the center of the mass of the maxilla. The abrasion measures 12 cm long by 2 cm wide and is approximately 1 cm deep at the center of the scar. There is evidence of new bone growth, especially near the ventral end of the gouge, demonstrating that this mark was made some days or weeks before death.</paragraph>
<paragraph id="8BB41B60E370D458FF378D44FD711169" blockId="2.[171,1415,183,2728]" pageId="2" pageNumber="235">Several perforations mar the anterior palatal shelf of the right maxilla. It could be that these are a result of postmortem weathering or a fungal infection. They do not seem to have been caused by osteomyelitis because cancellous bone is not evident.</paragraph>
<paragraph id="8BB41B60E370D458FF358C18FD9C12D1" blockId="2.[171,1415,183,2728]" pageId="2" pageNumber="235">
The nasal-maxillary suture is highly scalloped on 
<materialsCitation id="3B63113DE370D458FBF98C18FB13115E" ID-GBIF-Occurrence="3396384325" box="[1076,1235,1306,1342]" collectionCode="BHI" pageId="2" pageNumber="235" specimenCode="BHI 3033">BHI 3033</materialsCitation>
. Interlocking fingers of bone prevent any anterioposterior movement along this articulation. There is, however, the possibility of lateral movement, with the nasal-maxillary suture acting as a hinge, allowing the ventral portion of the maxilla to swing outward. If such movement were to occur, the interlocking ridges of the maxilla-maxilla palatal contact would have spread apart. Such kinetic movement would enhance the shock-absorbing abilities of the maxillae and teeth.
</paragraph>
<paragraph id="8BB41B60E370D458FF378FC1FD041330" blockId="2.[171,1415,183,2728]" pageId="2" pageNumber="235">Movement is also possible at the jugal-maxillary suture. Here the bones are joined by a tongue (jugal) and groove (maxilla) joint. This joint allows anterioposterior slippage.</paragraph>
<paragraph id="8BB41B60E370D458FF378E6BFED11D50" blockId="2.[171,1415,183,2728]" pageId="2" pageNumber="235">
<emphasis id="B97FC772E370D458FF378E6BFE1D13E6" box="[250,477,1897,1926]" italics="true" pageId="2" pageNumber="235">Lachrymals</emphasis>
: One of the characters uniting 
<taxonomicName id="4C0B60E3E370D458FC238E60FB2213E6" authorityName="Osborn" authorityYear="1905" box="[1006,1250,1890,1926]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
with 
<emphasis id="B97FC772E370D458FA8F8E60FE8713DD" italics="true" pageId="2" pageNumber="235">
<taxonomicName id="4C0B60E3E370D458FA8F8E60FE8113DD" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
,
</emphasis>
but separating it from other tyrannosaurs, such as 
<emphasis id="B97FC772E370D458FB608E9AFE0E1391" italics="true" pageId="2" pageNumber="235">
<taxonomicName id="4C0B60E3E370D458FB608E9AFF231391" baseAuthorityName="Currie" baseAuthorityYear="2003" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
, 
<taxonomicName id="4C0B60E3E370D458FF398ECFFE081391" authorityName="Cope" authorityYear="1866" box="[244,456,1997,2033]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
,
</emphasis>
and 
<emphasis id="B97FC772E370D458FDD28ECEFCCE1390" box="[543,782,1996,2032]" italics="true" pageId="2" pageNumber="235">
<taxonomicName id="4C0B60E3E370D458FDD28ECEFCC81390" authorityName="Osborn" authorityYear="1905" box="[543,776,1996,2032]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
,
</emphasis>
is the absence of a corneal process on the lachrymal (
<bibRefCitation id="EF9A6691E370D458FE638101FD981C46" author="Carr, T. D." box="[430,600,2050,2087]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="235" pagination="497 - 520" part="19" refId="ref5496" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19 (3): 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr 1999</bibRefCitation>
; 
<bibRefCitation id="EF9A6691E370D458FDA48100FBED1C46" author="Carr, T. D. &amp; Williamson, T. E." box="[617,1069,2050,2086]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="2" pageNumber="235" pagination="479 - 523" part="142" refId="ref5527" refString="Carr, T. D., and Williamson, T. E. 2004. Diversity of Late Maastrichtian Tryrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 142: 479 - 523." title="Diversity of Late Maastrichtian Tryrannosauridae (Dinosauria: Theropoda) from western North America" type="journal article" year="2004">Carr and Williamson 2004</bibRefCitation>
). This character also separates 
<taxonomicName id="4C0B60E3E370D458FE98813BFD871C3D" authorityName="Osborn" authorityYear="1905" box="[341,583,2105,2141]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
from 
<taxonomicName id="4C0B60E3E370D458FD75813AFC661C3C" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[696,934,2104,2140]" pageId="2" pageNumber="235">Nanotyrannus</taxonomicName>
(Larson in press). An additional character is the shape of the lachrymal: it is an inverted L shape in 
<taxonomicName id="4C0B60E3E370D458FF6381A1FE641CA7" authorityName="Osborn" authorityYear="1905" box="[174,420,2211,2247]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
, as it is in 
<emphasis id="B97FC772E370D458FD8181A0FCDE1CA6" box="[588,798,2210,2246]" italics="true" pageId="2" pageNumber="235">
<taxonomicName id="4C0B60E3E370D458FD8181A0FCD81CA6" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" box="[588,792,2210,2246]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
,
</emphasis>
and T shaped in 
<emphasis id="B97FC772E370D458FBF381A0FD931C9B" italics="true" pageId="2" pageNumber="235">
<taxonomicName id="4C0B60E3E370D458FBF381A0FAE71CA6" authorityName="Osborn" authorityYear="1905" box="[1086,1319,2210,2246]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
, 
<taxonomicName id="4C0B60E3E370D458FAF781A0FE811C9C" authorityName="Cope" authorityYear="1866" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
, 
<taxonomicName id="4C0B60E3E370D458FE9981D5FD931C9B" authority="(Carr et al. 2005)" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[340,595,2263,2299]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="235" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
</emphasis>
(
<bibRefCitation id="EF9A6691E370D458FDA181D5FCB51C9B" author="Carr, T. D. &amp; Williamson, T. E. &amp; Schwimrner, D. R." box="[620,885,2263,2299]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="235" pagination="119 - 143" part="25" refId="ref5570" refString="Carr, T. D., Williamson, T. E., and Schwimrner, D. R. 2005. A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama. Journal of Vertebrate Paleontology 25 (1): 119 - 143." title="A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama" type="journal article" year="2005">Carr et al. 2005</bibRefCitation>
), and 
<taxonomicName id="4C0B60E3E370D458FC1681D5FB091C9B" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[987,1225,2263,2299]" pageId="2" pageNumber="235">Nanotyrannus</taxonomicName>
(Larson in press).
</paragraph>
<paragraph id="8BB41B60E370D459FF348041FB3B151F" blockId="2.[171,1415,183,2728]" lastBlockId="3.[654,1899,183,2726]" lastPageId="3" lastPageNumber="236" pageId="2" pageNumber="235">
In addition to the pneumatic “lateral foramen” mentioned by 
<bibRefCitation id="EF9A6691E370D458FACB8040FED51DFC" author="Molnar, R. E." journalOrPublisher="Palaeontographica Abteilung A" pageId="2" pageNumber="235" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
, the lachrymals of 
<materialsCitation id="3B63113DE370D458FD918075FD3A1DFB" ID-GBIF-Occurrence="3396384352" box="[604,762,2423,2459]" collectionCode="BHI" pageId="2" pageNumber="235" specimenCode="BHI 3033">BHI 3033</materialsCitation>
possess 3 additional pneumatic openings. The first of these pneumatopores lies anterior to the lateral foramen, near the front of the lachrymal. It is on the dorsal margin of the antorbital fenestra and opens ventrally. In medial view, the lachrymals of 
<materialsCitation id="3B63113DE370D458FB2E8315FA401E5B" ID-GBIF-Occurrence="3396384322" box="[1251,1408,2583,2619]" collectionCode="BHI" pageId="2" pageNumber="235" specimenCode="BHI 3033">BHI 3033</materialsCitation>
expose 2 more pneumatic foramina. The medial lachrymal pneumatopore (Larson in press) opens anterior to and at the dorsal margin of a thin ridge that descends diagonally across the vertical ramus. This ridge terminates at the ventral, anterior border of the vertical ramus. Finally, a fourth pneumatopore (also medial) may be found at the front of the horizontal ramus. This pneumatopore opens anteriorly.
</paragraph>
<paragraph id="8BB41B60E371D459FD178892FAEC173D" blockId="3.[654,1899,183,2726]" pageId="3" pageNumber="236">
The lachrymals barely touch the ascending ramus of the maxilla, thus allowing movement. However, the nasal-lachrymal articulation locks quite firmly, with a somewhat scalloped surface on the medial aspect of the horizontal ramus of the lachrymal and a cleft on the lateral aspect of the horizontal ramus that receives the lachrymal process of the nasal (
<bibRefCitation id="EF9A6691E371D459F9538B67FCA316DD" author="Hurum, J. H. &amp; Sabath, K." journalOrPublisher="Acta Palaeontologia Polonica" pageId="3" pageNumber="236" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath 2003</bibRefCitation>
). The posterior surface at the junction of the horizontal and vertical rami is somewhat ball shaped. This ball inserts into a socket in the anterior surface of the frontals, suggesting possible prokinesis, or lifting of the muzzle (
<bibRefCitation id="EF9A6691E371D459FCA88A3BFADD173D" author="Larson P. L. &amp; Donnan, K." box="[869,1309,825,861]" journalOrPublisher="Invisible Cities Press, Montpelier, VT" pageId="3" pageNumber="236" refId="ref6014" refString="Larson P. L., and Donnan, K. 2002. Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life. Invisible Cities Press, Montpelier, VT" title="Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life" type="book" year="2002">Larson and Donnan 2002</bibRefCitation>
).
</paragraph>
<paragraph id="8BB41B60E371D459FD168A76FB1A10B2" blockId="3.[654,1899,183,2726]" pageId="3" pageNumber="236">
Nasals: As is the case with all tyrannosaurids, the nasals of 
<materialsCitation id="3B63113DE371D459F90A8A6DF8A217F3" ID-GBIF-Occurrence="3396384316" box="[1735,1890,879,915]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 3033">BHI 3033</materialsCitation>
are fused. This fusion, as well as lateral arching and the consistent thickness of the bone (2 cm or more over nearly the entire length of the nasals), provides an extremely strong structure for the dissemination of stress developed during feeding (see 
<bibRefCitation id="EF9A6691E371D459FBBA8D40FAA51006" author="Rayfield, E. J." box="[1143,1381,1090,1126]" journalOrPublisher="Proceedings of the Royal Society of London B" pageId="3" pageNumber="236" pagination="1451 - 1459" part="271" refId="ref6192" refString="Rayfield, E. J. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London B 271: 1451 - 1459" title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">Rayfield 2004</bibRefCitation>
). The nasal-frontal suture is a tongue (frontal) and groove (nasal) joint that allows fore-and-aft movement and would not preclude prokinesis.
</paragraph>
<paragraph id="8BB41B60E371D459FD118DEAFB6A11BD" blockId="3.[654,1899,183,2726]" pageId="3" pageNumber="236">
Postorbitals: The medial aspects of the postorbitals show a shallowcentral depression, or fossa. Other 
<taxonomicName id="4C0B60E3E371D459FB278C1AFA1E115C" authorityName="Osborn" authorityYear="1905" box="[1258,1502,1304,1340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="236" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
specimens (
<materialsCitation id="3B63113DE371D459F97A8C1AF89E115C" ID-GBIF-Occurrence="3396384358" box="[1719,1886,1304,1340]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 4812">BHI 4812</materialsCitation>
, 
<materialsCitation id="3B63113DE371D459FD5F8C4FFCFA1111" ID-GBIF-Occurrence="3396384349" box="[658,826,1357,1393]" collectionCode="MOR" pageId="3" pageNumber="236" specimenCode="MOR 555">MOR 555</materialsCitation>
, and 
<materialsCitation id="3B63113DE371D459FC5B8C4FFB801111" ID-GBIF-Occurrence="3396384354" box="[918,1088,1357,1393]" collectionCode="MOR" pageId="3" pageNumber="236" specimenCode="MOR 980">MOR 980</materialsCitation>
) also show this feature. Much deeper fossae are seen in 
<taxonomicName id="4C0B60E3E371D459FCDE8C80FC2611C6" authorityName="Cope" authorityYear="1866" box="[787,998,1410,1446]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="236" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E371D459FCDE8C80FC2611C6" box="[787,998,1410,1446]" italics="true" pageId="3" pageNumber="236">Gorgosaurus</emphasis>
</taxonomicName>
(TCM 2001.89.1), 
<taxonomicName id="4C0B60E3E371D459FAEA8C80F8D711C6" authority="(Currie 2003)" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[1319,1815,1410,1446]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="236" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E371D459FAEA8C80F9E611C6" box="[1319,1574,1410,1446]" italics="true" pageId="3" pageNumber="236">Daspletosaurus</emphasis>
(
<bibRefCitation id="EF9A6691E371D459F9F38C80F8CC11C6" author="Currie, P. J." box="[1598,1804,1410,1446]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="3" pageNumber="236" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
)
</taxonomicName>
, and 
<taxonomicName id="4C0B60E3E371D459FD5D8CBBFC4011BD" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[656,896,1465,1501]" pageId="3" pageNumber="236">Nanotyrannus</taxonomicName>
(Larson in press).
</paragraph>
<paragraph id="8BB41B60E371D459FD168CEFFA4C13E6" blockId="3.[654,1899,183,2726]" pageId="3" pageNumber="236">
Two paired, isolated osteoderms were found associated with the skull of 
<materialsCitation id="3B63113DE371D459FD718F20FC991226" ID-GBIF-Occurrence="3396384361" box="[700,857,1570,1606]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 3033">BHI 3033</materialsCitation>
(
<bibRefCitation id="EF9A6691E371D459FCBE8F20FB631226" author="Larson, N. L. &amp; Farrar, R. A. &amp; Shaffer, L." box="[883,1187,1570,1606]" editor="Wolberg, D. L. &amp; Stump, E. &amp; Rosenberg, G. D." journalOrPublisher="Academy of Natural Sciences, Philadelphia" pageId="3" pageNumber="236" pagination="68 - 69" refId="ref5861" refString="Larson, N. L., Farrar, R. A., and Shaffer, L. 1998. New information on the osteology of the skull of Tyrannosaurus rex. P. 68 - 69 in Wolberg, D. L., Stump, E., and Rosenberg, G. D. Dinofest International Proceedings. Academy of Natural Sciences, Philadelphia" title="New information on the osteology of the skull of Tyrannosaurus rex" type="proceedings paper" volumeTitle="Dinofest International Proceedings" year="1998">Larson et al. 1998</bibRefCitation>
). These dermal elements are analogous to the “postorbital rugosity” noted as fused to the postorbitals of some robust 
<taxonomicName id="4C0B60E3E371D459FD5C8F8FFC4412D1" authorityName="Osborn" authorityYear="1905" box="[657,900,1677,1713]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="236" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
specimens by 
<bibRefCitation id="EF9A6691E371D459FB468F8FFABE12D1" author="Molnar, R. E." box="[1163,1406,1677,1713]" journalOrPublisher="Palaeontographica Abteilung A" pageId="3" pageNumber="236" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
, 
<bibRefCitation id="EF9A6691E371D459FA5A8F8FF94612D1" author="Larson P. L." box="[1431,1670,1677,1713]" editor="Rosenberg, G. D. &amp; Wolberg, D. L." journalOrPublisher="Dino Fest. Paleontological Society Special Publication" pageId="3" pageNumber="236" pagination="139 - 155" refId="ref5930" refString="Larson P. L. 1994. Tyrannosaurus sex. P. 139 - 155 in Rosenberg, G. D., and Wolberg, D. L. (eds.). Dino Fest. Paleontological Society Special Publication No. 7" title="Tyrannosaurus sex" type="book chapter" year="1994">Larson (1994)</bibRefCitation>
, and 
<bibRefCitation id="EF9A6691E371D459F9208F8FFD3D1286" author="Brochu, C. A." journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="3" pageNumber="236" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003)</bibRefCitation>
. 
<taxonomicName id="4C0B60E3E371D459FCC28FC0FBC31286" authorityName="Osborn" authorityYear="1905" box="[783,1027,1730,1766]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="236" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
specimens with fused postorbital rugosities include 
<materialsCitation id="3B63113DE371D459FD5E8FF5FCFF137B" ID-GBIF-Occurrence="3396384317" box="[659,831,1783,1819]" collectionCode="MOR" pageId="3" pageNumber="236" specimenCode="MOR 008">MOR 008</materialsCitation>
, 
<materialsCitation id="3B63113DE371D459FC9C8FF5FBA0137B" ID-GBIF-Occurrence="3396384346" box="[849,1120,1783,1819]" collectionCode="FMNH" pageId="3" pageNumber="236" specimenCode="FMNH PR2081">FMNHPR2081</materialsCitation>
, 
<materialsCitation id="3B63113DE371D459FBBF8FF5FAD1137B" ID-GBIF-Occurrence="3396384337" box="[1138,1297,1783,1819]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 4812">BHI 4182</materialsCitation>
, and 
<materialsCitation id="3B63113DE371D459FAA68FF5F8E0137B" ID-GBIF-Occurrence="3396384340" box="[1387,1824,1783,1819]" collectionCode="UWGM" pageId="3" pageNumber="236" specimenCode="UWGM 181 (NS 1565.26)">UWGM 181 (NS 1565.26)</materialsCitation>
. On 
<materialsCitation id="3B63113DE371D459FD5F8E2FFCF31331" ID-GBIF-Occurrence="3396384370" box="[658,819,1837,1873]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 3033">BHI 3033</materialsCitation>
, these loose postorbital rugosities articulate with the postorbitals on the lateral surface directly above the orbit.
</paragraph>
<paragraph id="8BB41B60E371D459FD118E95F8E51D50" blockId="3.[654,1899,183,2726]" pageId="3" pageNumber="236">
The anterior surface of the postorbitals (above the orbit) and the posterior surface of the horizontal ramus of the lachrymals articulate with pyramidally shaped osteoderms, or horns. These horns were found as isolated elements in 
<materialsCitation id="3B63113DE371D459FCF68135FC211C3B" ID-GBIF-Occurrence="3396384330" box="[827,993,2103,2139]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 3033">BHI 3033</materialsCitation>
. They were noted by 
<bibRefCitation id="EF9A6691E371D459FAA88135F9961C3B" author="Brochu, C. A." box="[1381,1622,2103,2139]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="3" pageNumber="236" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003)</bibRefCitation>
as fused to the postorbitals in 
<materialsCitation id="3B63113DE371D459FC4D816EFB451CF0" ID-GBIF-Occurrence="3396384312" box="[896,1157,2156,2192]" collectionCode="FMNH" pageId="3" pageNumber="236" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
and were thought to be part of the postorbital rugosity of 
<bibRefCitation id="EF9A6691E371D459FC8081A3FBF81CA5" author="Molnar, R. E." box="[845,1080,2209,2245]" journalOrPublisher="Palaeontographica Abteilung A" pageId="3" pageNumber="236" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
. It is clear from 
<materialsCitation id="3B63113DE371D459FA8181A3FA281CA5" ID-GBIF-Occurrence="3396384375" box="[1356,1512,2209,2245]" collectionCode="BHI" pageId="3" pageNumber="236" specimenCode="BHI 3033">BHI 3033</materialsCitation>
(a set of isolated horns was also found with 
<materialsCitation id="3B63113DE371D459FC1781D5FB4E1C9B" ID-GBIF-Occurrence="3396384315" box="[986,1166,2263,2299]" collectionCode="MOR" pageId="3" pageNumber="236" specimenCode="MOR 1125">MOR 1125</materialsCitation>
) that they are independent dermal elements. These horns bridge the gap between the postorbital and the lachrymal.
</paragraph>
<paragraph id="8BB41B60E371D459FD108043FA491E5A" blockId="3.[654,1899,183,2726]" pageId="3" pageNumber="236">The postorbital-frontal suture is interfingered and allows no movement. The joint between the postorbital and the jugal is smooth and planar, allowing diagonal slippage. The postorbital-squamosal joint is an expanding tongue and groove, which allows fore-and-aft separation, although kinesis at this point is difficult to reconstruct.</paragraph>
<paragraph id="8BB41B60E371D45EFD128349FD811739" blockId="3.[654,1899,183,2726]" lastBlockId="4.[170,1411,176,2668]" lastPageId="4" lastPageNumber="237" pageId="3" pageNumber="236">
SQUAMOSALS: The central portions of the anterior aspect of the squamosals are perforated by a large pneumatopore (most clearly seen in 
<materialsCitation id="3B63113DE371D45EF8EE8382FF3D14BB" ID-GBIF-Occurrence="3396384311" collectionCode="BHI" lastPageId="4" lastPageNumber="237" pageId="3" pageNumber="236" specimenCode="BHI 4100">BHI 4100</materialsCitation>
). This pneumatopore is also present in 
<taxonomicName id="4C0B60E3E376D45EFC5889B5FBA114BB" authorityName="Maleev" authorityYear="1955" box="[917,1121,183,219]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="237" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E376D45EFC5889B5FBA114BB" box="[917,1121,183,219]" italics="true" pageId="4" pageNumber="237">Tarbosaurus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9A6691E376D45EFBB489B5FE90156F" author="Hurum, J. H. &amp; Sabath, K." journalOrPublisher="Acta Palaeontologia Polonica" pageId="4" pageNumber="237" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath 2003</bibRefCitation>
), but it is absent in 
<emphasis id="B97FC772E376D45EFD4689E9FCA3156F" box="[651,867,235,271]" italics="true" pageId="4" pageNumber="237">
<taxonomicName id="4C0B60E3E376D45EFD4689E9FC9D156F" authority=", Nanotyrannus" authorityName="Cope" authorityYear="1866" box="[651,861,235,271]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="237" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
,
</emphasis>
<taxonomicName id="4C0B60E3E376D45EFCA289E9FB9C156F" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[879,1116,235,271]" pageId="4" pageNumber="237">Nanotyrannus</taxonomicName>
(Larson in press), and 
<taxonomicName id="4C0B60E3E376D45EFF3D8823FD1B1525" authority="(Currie 2003)" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[240,731,289,325]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="237" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E376D45EFF3D8823FE2F1525" box="[240,495,289,325]" italics="true" pageId="4" pageNumber="237">Daspletosaurus</emphasis>
(
<bibRefCitation id="EF9A6691E376D45EFDC88823FD101525" author="Currie, P. J." box="[517,720,289,325]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="4" pageNumber="237" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
)
</taxonomicName>
. Just anterior of center, the lateral ventral borders of the V-shaped squamosals are marked by a deep fold. This fold corresponds to a deep concavity on the squamosal process noted by 
<bibRefCitation id="EF9A6691E376D45EFAC98889FECC1585" author="Molnar, R. E." journalOrPublisher="Palaeontographica Abteilung A" pageId="4" pageNumber="237" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
on the quadratojugal. These 2 surfaces could have been the origin and insertion for a ligament connecting the 2 elements (see Quadratojugals, below). The articulation of the squamosal with the exoccipital is fairly flat and may have allowed limited sliding motion. The squamosal-quadrate joint is well developed and is a double ball-in-socket (quadrate-in-squamosal) joint. The 2 balls are side by side and are connected by a saddle that is reversed in the squamosal. This joint gives great flexibility fore and aft, but limits lateral movement.
</paragraph>
<paragraph id="8BB41B60E376D45EFF388A6CFD31126C" blockId="4.[170,1411,176,2668]" pageId="4" pageNumber="237">
JUGALS: Both jugals have pathological, healed puncture wounds (Larson 2001). These may have been the result of face-biting behavior described by 
<bibRefCitation id="EF9A6691E376D45EFF148AD7FDAD1799" author="Tanke, D. H. &amp; Currie, P. J." box="[217,621,981,1017]" editor="Perez-Moreno, B. P &amp; Holtz, T. J. &amp; Sanz, J. L. &amp; Moratalla, J." journalOrPublisher="Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon" pageId="4" pageNumber="237" pagination="167 - 184" part="15" refId="ref6301" refString="Tanke, D. H., and Currie, P. J. 1998. Elead-biting behavior in theropod dinosaurs: paleopathological evidence. P. 167 - 184 in Perez-Moreno, B. P, Holtz, T. J., Sanz, J. L., and Moratalla, J. (eds.). Aspects ofTheropod Paleobiology. Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon, 15." title="Elead-biting behavior in theropod dinosaurs: paleopathological evidence" type="book" volumeTitle="Aspects ofTheropod Paleobiology" year="1998">Tanke and Currie (1998)</bibRefCitation>
. The injury to the left jugal is a circular hole, 3.5 cm by 2.5 cm, that penetrates at a point near the origin of the cleft that receives the quadratojugal (here the jugal is approximately 4 cm thick). In medial aspect, the perforation emerges, leaving an opening 2.7 cm by 1.5 cm. The injury to the right jugal is located just anterior of center of the ascending ramus, at the ventral termination of the postorbital-jugal joint. The lateral opening measures 2 cm by 2.5 cm. The medial exit is larger (3.5 cm by 6.5 cm) and somewhat triangular in shape. 
<bibRefCitation id="EF9A6691E376D45EFC8C8C4BFBEA110D" author="Molnar, R. E." box="[833,1066,1353,1389]" journalOrPublisher="Palaeontographica Abteilung A" pageId="4" pageNumber="237" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
showed that there is pneumatic sinus in this area (also seen in 
<materialsCitation id="3B63113DE376D45EFC868C7CFC2511C2" ID-GBIF-Occurrence="3396384372" box="[843,997,1406,1442]" collectionCode="BHI" pageId="4" pageNumber="237" specimenCode="BHI 3033">BHI 3033</materialsCitation>
) that separates the lateral layer from the medial layer of bone. The larger exit hole may be explained by the thinness of that medial layer.
</paragraph>
<paragraph id="8BB41B60E376D45EFF388F1FFA8B13E1" blockId="4.[170,1411,176,2668]" pageId="4" pageNumber="237">The jugal-ectopterygoid joint is smooth, allowing palatal kinesis (see Larsson this volume). The articulation of the jugal and the quadratojugal is also smooth, marked by the cleft mentioned above. A horizontal ridge on the lateral aspect of the jugal process of the quadratojugal fits into this cleft. Fore-and-aft movement of the quadratojugal along this joint, with the firmly attached quadrate (see below) rocking on the double ball-and-socket squamosal-quadrate joint described above, would produce streptostyly.</paragraph>
<paragraph id="8BB41B60E376D45EFF3B8E95FD4F1DF6" blockId="4.[170,1411,176,2668]" pageId="4" pageNumber="237">
Quadratojugals: The left quadratojugal is isolated, and the right one is firmly attached to the quadrate. Although 
<bibRefCitation id="EF9A6691E376D45EFBC18EC5FB35138B" author="Molnar, R. E." box="[1036,1269,1991,2027]" journalOrPublisher="Palaeontographica Abteilung A" pageId="4" pageNumber="237" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991</bibRefCitation>
, p. 161) presumed that “no articulation with the squamosal existed,” there might have been a ligamental attachment (see Squamosals, above) that joined the 2 elements. This ligament could have acted as a tensile spring, returning the quadrate-quadratojugal to rest position after a streptostylic extension. The quadratojugal-quadrate articulation is somewhat edentate, creating a nonmovable joint. This suture also accounts for the firm attachment of the right quadratojugal to the right quadrate in 
<materialsCitation id="3B63113DE376D45EFC46803EFBE81D00" ID-GBIF-Occurrence="3396384359" box="[907,1064,2364,2400]" collectionCode="BHI" pageId="4" pageNumber="237" specimenCode="BHI 3033">BHI 3033</materialsCitation>
and other disarticulated skulls (i.e., 
<materialsCitation id="3B63113DE376D45EFE0E8070FDBF1DF6" ID-GBIF-Occurrence="3396384334" box="[451,639,2418,2454]" collectionCode="MOR" pageId="4" pageNumber="237" specimenCode="MOR 1125">MOR 1125</materialsCitation>
).
</paragraph>
<paragraph id="8BB41B60E376D45FFF3A80AEFA90156A" blockId="4.[170,1411,176,2668]" lastBlockId="5.[654,1897,173,961]" lastPageId="5" lastPageNumber="238" pageId="4" pageNumber="237">
Quadrates: The quadrates of 
<materialsCitation id="3B63113DE376D45EFCDD80A5FC721DAB" ID-GBIF-Occurrence="3396384347" box="[784,946,2471,2507]" collectionCode="BHI" pageId="4" pageNumber="237" specimenCode="BHI 3033">BHI 3033</materialsCitation>
are both preserved with a small notch (1 cm in diameter) on the dorsal edge of the squamosal process. This notch lies just anterior to the articulation with the quadrate. The notch probably corresponds to a much larger notch (several centimeters in diameter) found in 
<emphasis id="B97FC772E377D45FFC1B89B0FB7014B6" box="[982,1200,178,214]" italics="true" pageId="5" pageNumber="238">
<taxonomicName id="4C0B60E3E377D45FFC1B89B0FB6A14B6" authority=", Nanotyrannus" authorityName="Cope" authorityYear="1866" box="[982,1194,178,214]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
,
</emphasis>
<taxonomicName id="4C0B60E3E377D45FFB7389B0FA6C14B6" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[1214,1452,178,214]" pageId="5" pageNumber="238">Nanotyrannus</taxonomicName>
(Larson in press), 
<emphasis id="B97FC772E377D45FF91289B0FCC2156A" italics="true" pageId="5" pageNumber="238">
<taxonomicName id="4C0B60E3E377D45FF91289B0FD3C156A" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
,
</emphasis>
and 
<taxonomicName id="4C0B60E3E377D45FFC9489E4FA8B156A" authority="(Currie 2003)" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[857,1355,230,266]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E377D45FFC9489E4FB9A156A" box="[857,1114,230,266]" italics="true" pageId="5" pageNumber="238">Daspletosaurus</emphasis>
(
<bibRefCitation id="EF9A6691E377D45FFBBC89E4FA80156A" author="Currie, P. J." box="[1137,1344,230,266]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="5" pageNumber="238" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BB41B60E377D45FFD14881FFA2716E0" blockId="5.[654,1897,173,961]" pageId="5" pageNumber="238">
The quadrate-pterygoid joint surfaces are smooth, allowing streptostyly (hinted at by 
<bibRefCitation id="EF9A6691E377D45FFC078850FB671516" author="Molnar, R. E." box="[970,1191,338,374]" journalOrPublisher="Palaeontographica Abteilung A" pageId="5" pageNumber="238" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar 1991</bibRefCitation>
).The proximal end has a double articular surface (see Squamosals, above). This is similar to the situation in 
<taxonomicName id="4C0B60E3E377D45FF9388884FD3A1580" authorityName="Cope" authorityYear="1866" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E377D45FF9388884FD3A1580" italics="true" pageId="5" pageNumber="238">Gorgosaurus</emphasis>
</taxonomicName>
(TCM 2001.89.1) and 
<taxonomicName id="4C0B60E3E377D45FFBBB88BEF9A31580" authority="(Currie 2003)" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[1142,1635,444,480]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E377D45FFBBB88BEFAB51580" box="[1142,1397,444,480]" italics="true" pageId="5" pageNumber="238">Daspletosaurus</emphasis>
(
<bibRefCitation id="EF9A6691E377D45FFA4188BEF9981580" author="Currie, P. J." box="[1420,1624,444,480]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="5" pageNumber="238" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
)
</taxonomicName>
, whereas 
<taxonomicName id="4C0B60E3E377D45FF8CD88BEFCD91674" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E377D45FF8CD88BEFCD91674" italics="true" pageId="5" pageNumber="238">Albertosaurus</emphasis>
</taxonomicName>
(BHI 6234) and 
<taxonomicName id="4C0B60E3E377D45FFB8E88F2FAF31674" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[1091,1331,496,532]" pageId="5" pageNumber="238">Nanotyrannus</taxonomicName>
have only a single ball-in-socket articulation. Although all tyrannosaurs exhibit the capacity for streptostyly, it seems that it was accomplished in different ways.
</paragraph>
<paragraph id="8BB41B60E377D45FFD178B93F8D917DE" blockId="5.[654,1897,173,961]" pageId="5" pageNumber="238">
The double condylar surface of the quadrates at the quadrate-articular joint is oriented in such a way that it forms a kind of screw on the surface of the main joint for the opening and closing of the jaws. As the jaws opened, this screw would, in effect, widen the gape of the jaws by forcing the articulars away from each other. Likewise, this screw would bring the rear of the jaws back together again as the jaws closed (
<bibRefCitation id="EF9A6691E377D45FF9E28A98F8C917DE" author="Molnar, R. E." box="[1583,1801,922,958]" journalOrPublisher="Palaeontographica Abteilung A" pageId="5" pageNumber="238" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar 1991</bibRefCitation>
).
</paragraph>
<paragraph id="8BB41B60E377D45FFF678D3AFE2F100A" blockId="5.[170,495,1080,1130]" box="[170,495,1080,1130]" pageId="5" pageNumber="238">
<heading id="D0FCAC0CE377D45FFF678D3AFE2F100A" bold="true" box="[170,495,1080,1130]" centered="true" fontSize="15" level="2" pageId="5" pageNumber="238" reason="0">
<emphasis id="B97FC772E377D45FFF678D3AFEEF100A" bold="true" box="[170,303,1080,1130]" pageId="5" pageNumber="238">Palatal</emphasis>
Complex
</heading>
</paragraph>
<paragraph id="8BB41B60E377D45FFD118D47FC46132D" blockId="5.[656,1898,1084,2722]" pageId="5" pageNumber="238">Pterygoids: Both pterygoids are complete. The articulation with the quadrate is discussed above. The rear of the palatal plate wraps over the basisphenoid, limiting posterior movement. Although the dorsal surface of the palatal plate of the pterygoid is fairly smooth, little movement probably occurred at the ectopterygoid-ptervgoid joint, and the folding of the quadrate process of the pterygoid blocked the ectopterygoid from moving posteriorly relative to the pterygoid. The folded posterior pterygoid-palatine joint allowed fore-and-aft movement but restricted lateral movement. The smooth anterior pterygoid-palatine joints allow fore-and-aft movement. The vomerine process of the pterygoid is a smooth vertical blade that touches the palatine on its lateral surface and the vomer on its medial surface The pterygoid-vomer suture is bounded by lateral extensions of the bifurcate stem. These 1-cm-deep lips limit forward movement of the pterygoid relative to the vomer. See Larsson (this volume) for a discussion of palatal kinesis.</paragraph>
<paragraph id="8BB41B60E377D45FFD108E66FCFF1E62" blockId="5.[656,1898,1084,2722]" pageId="5" pageNumber="238">
Epipterygoids: The epipterygoids articulate on the anterior surface of the vertical quadrate process of the pterygoids. The epipterygoids of 
<materialsCitation id="3B63113DE377D45FF8EC8E96FD1D138D" ID-GBIF-Occurrence="3396384320" collectionCode="BHI" pageId="5" pageNumber="238" specimenCode="BHI 3033">BHI 3033</materialsCitation>
are similar to those described for 
<taxonomicName id="4C0B60E3E377D45FFAD18ECBF8CE138D" authority="(Currie 2003)" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[1308,1806,1993,2029]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E377D45FFAD18ECBF9DC138D" box="[1308,1564,1993,2029]" italics="true" pageId="5" pageNumber="238">Daspletosaurus</emphasis>
(
<bibRefCitation id="EF9A6691E377D45FF9F88ECBF8C3138D" author="Currie, P. J." box="[1589,1795,1993,2029]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="5" pageNumber="238" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
)
</taxonomicName>
. The angle of the inverted V in 
<materialsCitation id="3B63113DE377D45FFB9D8EFFFB2D1C41" ID-GBIF-Occurrence="3396384353" box="[1104,1261,2045,2081]" collectionCode="BHI" pageId="5" pageNumber="238" specimenCode="BHI 3033">BHI 3033</materialsCitation>
is, however, slightly greater than that of 
<emphasis id="B97FC772E377D45FFD0D8136FC0C1C38" box="[704,972,2100,2136]" italics="true" pageId="5" pageNumber="238">Dasplatosaurus,</emphasis>
and the concavity at the base, mentioned by 
<bibRefCitation id="EF9A6691E377D45FF93A8136FD3B1CED" author="Currie, P. J." journalOrPublisher="Acta Palaeontologia Polonica" pageId="5" pageNumber="238" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie (2003)</bibRefCitation>
, is much deeper in 
<taxonomicName id="4C0B60E3E377D45FFB99816BFA431CED" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1108,1411,2153,2189]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="238" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
to such an extent that the 2 arms of the 
<emphasis id="B97FC772E377D45FFC92819CFCBD1CA2" bold="true" box="[863,893,2206,2242]" pageId="5" pageNumber="238">V</emphasis>
are actually bifurcated. The more medial wing of the epipterygoid butts against the anterior medial edge of the quadrate process of the pterygoid, continuing the curve, established by the medial edge of the quadrate process. This butt joint restricts movement (other than folding) at the epipterygoid-pterygoid joint. The more dorsal epipterygoid-laterosphenoid articulations are smooth and would allow movement during palatal kinesis.
</paragraph>
<paragraph id="8BB41B60E377D45CFD108315FEBD1512" blockId="5.[656,1898,1084,2722]" lastBlockId="6.[164,1407,175,1594]" lastPageId="6" lastPageNumber="239" pageId="5" pageNumber="238">
Ectopterygoids: The hook-shaped ectopterygoids of 
<materialsCitation id="3B63113DE377D45FF94A8310F8E51E56" ID-GBIF-Occurrence="3396384302" box="[1671,1829,2578,2614]" collectionCode="BHI" pageId="5" pageNumber="238" specimenCode="BHI 3033">BHI 3033</materialsCitation>
are perforated with a large pneumatopore typical of theropods (
<bibRefCitation id="EF9A6691E377D45FF9B1834AF8951E0C" author="Molnar, R. E." box="[1660,1877,2632,2668]" journalOrPublisher="Palaeontographica Abteilung A" pageId="5" pageNumber="238" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar 1991</bibRefCitation>
). 
<bibRefCitation id="EF9A6691E377D45FFD5F837FFC751EC1" author="Carr, T. D. &amp; Williamson, T. E. &amp; Schwimrner, D. R." box="[658,949,2685,2721]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="5" pageNumber="238" pagination="119 - 143" part="25" refId="ref5570" refString="Carr, T. D., Williamson, T. E., and Schwimrner, D. R. 2005. A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama. Journal of Vertebrate Paleontology 25 (1): 119 - 143." title="A new genus of tyrannosaurid from the Late Cretaceous (Middle Campanian) Demopolis Formation of Alabama" type="journal article" year="2005">Carr et al. (2005)</bibRefCitation>
noted that the thick lip that bounds the pneumatopore was a character for 
<taxonomicName id="4C0B60E3E374D45CFE2689ADFD2214B3" authorityName="Osborn" authorityYear="1905" box="[491,738,175,211]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
. The anterior limb of the ectopterygoid contacts the ventral medial surface near the center of the jugal. The smooth ectopterygoid-jugal joint is critical for palatal kinesis (see Larsson this volume).
</paragraph>
<paragraph id="8BB41B60E374D45CFF3F888BFE6B1045" blockId="6.[164,1407,175,1594]" pageId="6" pageNumber="239">
Palatines: The palatine was incompletely described by 
<bibRefCitation id="EF9A6691E374D45CFB438886FAB515C8" author="Molnar, R. E." box="[1166,1397,388,424]" journalOrPublisher="Palaeontographica Abteilung A" pageId="6" pageNumber="239" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991)</bibRefCitation>
. Although both palatines are complete in 
<materialsCitation id="3B63113DE374D45CFCA688BAFBCF15BC" ID-GBIF-Occurrence="3396384309" box="[875,1039,440,476]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 3033">BHI 3033</materialsCitation>
, they were somewhat weathered before burial, making a complete assessment difficult. 
<materialsCitation id="3B63113DE374D45CFB1A88ECFAB51672" ID-GBIF-Occurrence="3396384368" box="[1239,1397,494,530]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 4100">BHI 4100</materialsCitation>
, however, does include an exceptionally well-preserved right palatine that aids in the interpretation of 
<materialsCitation id="3B63113DE374D45CFDBB8B5AFCD6161C" ID-GBIF-Occurrence="3396384335" box="[630,790,600,636]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 3033">BHI 3033</materialsCitation>
. This palatine has a large pneumatopore on the central lateral surface near the articulation with the maxilla, which opens into a vast chamber. There is also a deep pneumatic fossa directly anterior to this pneumatopore and of near-equal size. The palatines of 
<taxonomicName id="4C0B60E3E374D45CFF6A8A2CFE531732" authorityName="Osborn" authorityYear="1905" box="[167,403,814,850]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
are much more inflated than those of 
<taxonomicName id="4C0B60E3E374D45CFBCD8A2CFB3B1732" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[1024,1275,814,850]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E374D45CFBCD8A2CFB3B1732" box="[1024,1275,814,850]" italics="true" pageId="6" pageNumber="239">Daspletosaurus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9A6691E374D45CFAC28A2CFF3917E7" author="Currie, P. J." journalOrPublisher="Acta Palaeontologia Polonica" pageId="6" pageNumber="239" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
; Carr etal. 2005), 
<taxonomicName id="4C0B60E3E374D45CFDEE8A60FCC517E6" authorityName="Osborn" authorityYear="1905" box="[547,773,866,902]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E374D45CFDEE8A60FCC517E6" box="[547,773,866,902]" italics="true" pageId="6" pageNumber="239">Albertosaurus</emphasis>
</taxonomicName>
(Carr etal. 2005), 
<taxonomicName id="4C0B60E3E374D45CFBFA8A60FAC517E6" authorityName="Cope" authorityYear="1866" box="[1079,1285,866,902]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E374D45CFBFA8A60FAC517E6" box="[1079,1285,866,902]" italics="true" pageId="6" pageNumber="239">Gorgosaurus</emphasis>
</taxonomicName>
(TCM 2001.89.1), and 
<taxonomicName id="4C0B60E3E374D45CFE6E8A9AFD4C17DC" authority="Gilmore, 1946" authorityName="Gilmore" authorityYear="1946" box="[419,652,920,956]" pageId="6" pageNumber="239">Nanotyrannus</taxonomicName>
(BMRP 2002.4.1). However, their similarity to 
<taxonomicName id="4C0B60E3E374D45CFF6A8ACFFEAB1791" authorityName="Maleev" authorityYear="1955" box="[167,363,973,1009]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E374D45CFF6A8ACFFEAB1791" box="[167,363,973,1009]" italics="true" pageId="6" pageNumber="239">Tarbosaurus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9A6691E374D45CFE4F8ACFFCDA1791" author="Hurum, J. H. &amp; Sabath, K." box="[386,794,973,1009]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="6" pageNumber="239" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath 2003</bibRefCitation>
) could indicate that this inflation is a function of size.
</paragraph>
<paragraph id="8BB41B60E374D45CFF3C8D3AFCB81105" blockId="6.[164,1407,175,1594]" pageId="6" pageNumber="239">The palatine-maxilla joint is slightly grooved, with corresponding low tongues on the opposing bones. This joint would allow limited movement fore and aft and would not preclude lateral movement of the maxillae. The palatines also contact the anterior medial surface of the jugals and the ventral medial surface of the lachrymals in a smooth joint that would not restrict palatal kinesis or lateral movement.</paragraph>
<paragraph id="8BB41B60E374D45CFF3F8C7EFAAC125A" blockId="6.[164,1407,175,1594]" pageId="6" pageNumber="239">
<emphasis id="B97FC772E374D45CFF3F8C7EFEAD11F9" box="[242,365,1404,1433]" italics="true" pageId="6" pageNumber="239">Vomer</emphasis>
: The vomer of 
<materialsCitation id="3B63113DE374D45CFD598C75FCFA11FB" ID-GBIF-Occurrence="3396384364" box="[660,826,1399,1435]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 3033">BHI 3033</materialsCitation>
matches 
<bibRefCitation id="EF9A6691E374D45CFC3D8C75FB3C11FB" author="Molnar, R. E." box="[1008,1276,1399,1435]" journalOrPublisher="Palaeontographica Abteilung A" pageId="6" pageNumber="239" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar’s (1991)</bibRefCitation>
reconstructed description. The anterior rhomboid plate in 
<materialsCitation id="3B63113DE374D45CFBFE8CAEFB1211B0" ID-GBIF-Occurrence="3396384336" box="[1075,1234,1452,1488]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 3033">BHI 3033</materialsCitation>
is convex in dorsal aspect and concave ventrally. This rhomboid plate also sports a small healed puncture wound near the base (
<bibRefCitation id="EF9A6691E374D45CFC688F14FA9C125A" author="Larson P. L. &amp; Donnan, K." box="[933,1372,1558,1594]" journalOrPublisher="Invisible Cities Press, Montpelier, VT" pageId="6" pageNumber="239" refId="ref6014" refString="Larson P. L., and Donnan, K. 2002. Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life. Invisible Cities Press, Montpelier, VT" title="Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life" type="book" year="2002">Larson and Donnan 2002</bibRefCitation>
).
</paragraph>
<paragraph id="8BB41B60E374D45CFA068FB1F9541342" blockId="6.[1482,1884,1715,1826]" pageId="6" pageNumber="239">Braincase, Including Skull Roof</paragraph>
<paragraph id="8BB41B60E374D45CFF658FBEFC571C94" blockId="6.[167,1407,1724,2292]" pageId="6" pageNumber="239">
<materialsCitation id="3B63113DE374D45CFF658FBEFE811280" ID-GBIF-Occurrence="3396384338" box="[168,321,1724,1760]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 3033">BHI 3033</materialsCitation>
has a relatively complete braincase that is missing only the ventral portion of the basicranium (as a result of predepositional weathering). As in 
<materialsCitation id="3B63113DE374D45CFF658E24FE7A132A" ID-GBIF-Occurrence="3396384329" box="[168,442,1830,1866]" collectionCode="FMNH" pageId="6" pageNumber="239" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
, the prefrontals are firmly affixed to the frontals (
<bibRefCitation id="EF9A6691E374D45CFACC8E24FF3913E0" author="Brochu, C. A." journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="6" pageNumber="239" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu 2003</bibRefCitation>
). The subadult 
<materialsCitation id="3B63113DE374D45CFE3C8E5EFD5813E0" ID-GBIF-Occurrence="3396384363" box="[497,664,1884,1920]" collectionCode="MOR" pageId="6" pageNumber="239" specimenCode="MOR 1125">MOR 1125</materialsCitation>
has loose prefrontals, and a mature individual of 
<taxonomicName id="4C0B60E3E374D45CFF028E8FFEE513D5" authorityName="Osborn" authorityYear="1905" box="[207,293,1933,1973]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="239" phylum="Chordata" rank="species" species="rex">
T. 
<emphasis id="B97FC772E374D45CFF3A8E93FEE513D5" box="[247,293,1937,1973]" italics="true" pageId="6" pageNumber="239">rex</emphasis>
</taxonomicName>
A (see N. L. Larson this volume) preserves both frontals as disarticulated elements. A healed puncture wound is located near the center of the supraoccipital crest of the left parietal. This 3-cm-diameter hole perforates what was originally 3 cm of dense bone. Immediately above the perforation, a 12 cm by 5 cm fragment of bone is missing from the edge of the supraoccipital crest, presumably the result of the same incident causing the perforation (Larson 2001; 
<bibRefCitation id="EF9A6691E374D45CFE1081D2FC471C94" author="Larson P. L. &amp; Donnan, K." box="[477,903,2256,2292]" journalOrPublisher="Invisible Cities Press, Montpelier, VT" pageId="6" pageNumber="239" refId="ref6014" refString="Larson P. L., and Donnan, K. 2002. Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life. Invisible Cities Press, Montpelier, VT" title="Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life" type="book" year="2002">Larson and Donnan 2002</bibRefCitation>
).
</paragraph>
<paragraph id="8BB41B60E374D45CFA06806DF95A1DC1" blockId="6.[1483,1690,2415,2465]" box="[1483,1690,2415,2465]" pageId="6" pageNumber="239">Mandibles</paragraph>
<paragraph id="8BB41B60E374D45CFF68807EFD191EC5" blockId="6.[165,1406,2423,2725]" pageId="6" pageNumber="239">
Articular: Only the left articular was recovered from 
<materialsCitation id="3B63113DE374D45CFBAD8075FAC61DFB" ID-GBIF-Occurrence="3396384360" box="[1120,1286,2423,2459]" collectionCode="BHI" pageId="6" pageNumber="239" specimenCode="BHI 3033">BHI 3033</materialsCitation>
; it was tightly articulated to the left surangular. The articular surface of the exposed right articular-surangular suture shows a great deal of interfingering, eliminating the possibility of movement at this suture. Likewise, the nature of the articulation with the prearticulars reveals that the articular was firmly locked into position in life.
</paragraph>
<paragraph id="8BB41B60E374D45CFA0583F7F9C01F70" blockId="6.[1480,1536,2805,2832]" box="[1480,1536,2805,2832]" pageId="6" pageNumber="239">
<pageNumber id="BBA010B6E374D45CFA0583F7F9C01F70" ambiguity="4" box="[1480,1536,2805,2832]" fuzzyness="0" pageId="6" pageNumber="239" score="31.0" value="239">239</pageNumber>
</paragraph>
<paragraph id="8BB41B60E375D45DFD1389B2FCB015C3" blockId="7.[658,1904,165,2708]" pageId="7" pageNumber="240">Prearticulars: The prearticulars thin anteriorly to approximately 2 mm in lateral dimension for the final third of their length, although they measure an average of 8 cm in width. Both of the prearticulars show small (several millimeters), possibly pathologic perforations near their anterior terminations.</paragraph>
<paragraph id="8BB41B60E375D45DFD1188BBFC8E172C" blockId="7.[658,1904,165,2708]" pageId="7" pageNumber="240">
<emphasis id="B97FC772E375D45DFD1188BBFC5415B6" box="[732,916,441,470]" italics="true" pageId="7" pageNumber="240">Angulars</emphasis>
: The angulars of 
<materialsCitation id="3B63113DE375D45DFB7188B1FA9815B7" ID-GBIF-Occurrence="3396384356" box="[1212,1368,435,471]" collectionCode="BHI" pageId="7" pageNumber="240" specimenCode="BHI 3033">BHI 3033</materialsCitation>
match 
<bibRefCitation id="EF9A6691E375D45DFA1488B1F91815B7" author="Molnar, R. E." box="[1497,1752,435,471]" journalOrPublisher="Palaeontographica Abteilung A" pageId="7" pageNumber="240" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar’s (1991)</bibRefCitation>
description. The left angular shows some minor remodeling on the anterior most surface, corresponding to a puncture wound (where it articulates) on the left surangular. The spatulate posterior medial surface articulates with the surangular dorsally and the prearticular ventrally. Anteriorly, the angular thins dorsoventrally and is sandwiched between the dentary (laterally) and the splenial (medially). All these surfaces are smooth and allow fore-and-aft movement.
</paragraph>
<paragraph id="8BB41B60E375D45DFD128A63F98C11A9" blockId="7.[658,1904,165,2708]" pageId="7" pageNumber="240">
SURANGULARS: Both clam-shaped surangulars show healed puncture wounds. The right surangular has a large, remodeled perforation just anterior to and the same size as the surangular fenestra; a second perforation is on the anterior edge of this surangular. The left surangular is even more pathologic, with 4 large perforations. All show evidence of healing. They are rounded and thickened at the edges by remodeling. This is not unusual for 
<emphasis id="B97FC772E375D45DFD078D9EFC0610A0" box="[714,966,1180,1216]" italics="true" pageId="7" pageNumber="240">
<taxonomicName id="4C0B60E3E375D45DFD078D9EFC0010A0" authorityName="Osborn" authorityYear="1905" box="[714,960,1180,1216]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="240" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
,
</emphasis>
and perforations of this large, thin element are found in nearly every preserved surangular. 
<materialsCitation id="3B63113DE375D45DFB118DD3FA261095" ID-GBIF-Occurrence="3396384313" box="[1244,1510,1233,1269]" collectionCode="FMNH" pageId="7" pageNumber="240" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
(Larson 2001), 
<materialsCitation id="3B63113DE375D45DF93F8DD3FD3B1149" ID-GBIF-Occurrence="3396384371" collectionCode="LACM" pageId="7" pageNumber="240" specimenCode="LACM 23844">LACM 23844</materialsCitation>
(
<bibRefCitation id="EF9A6691E375D45DFCDF8C07FC2B1149" author="Molnar, R. E." box="[786,1003,1285,1321]" journalOrPublisher="Palaeontographica Abteilung A" pageId="7" pageNumber="240" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar 1991</bibRefCitation>
), 
<materialsCitation id="3B63113DE375D45DFBC88C07FB221149" ID-GBIF-Occurrence="3396384348" box="[1029,1250,1285,1321]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761" pageId="7" pageNumber="240" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
(Osborn 1916), 
<materialsCitation id="3B63113DE375D45DFA238C07F94E1149" ID-GBIF-Occurrence="3396384331" box="[1518,1678,1285,1321]" collectionCode="MOR" pageId="7" pageNumber="240" specimenCode="MOR 555">MOR 555</materialsCitation>
(
<bibRefCitation id="EF9A6691E375D45DF9658C07FCB41100" author="Horner, J. R. &amp; Lessem, D." journalOrPublisher="Simon &amp; Schuster, New York" pageId="7" pageNumber="240" refId="ref5791" refString="Horner, J. R., and Lessem, D. 1993. The Complete T. rex. Simon &amp; Schuster, New York." title="The Complete T. rex" type="book" year="1993">Horner and Lessem 1993</bibRefCitation>
), 
<materialsCitation id="3B63113DE375D45DFC5B8C3EFB871100" ID-GBIF-Occurrence="3396384373" box="[918,1095,1340,1376]" collectionCode="MOR" pageId="7" pageNumber="240" specimenCode="MOR 008">MOR 008</materialsCitation>
, 
<materialsCitation id="3B63113DE375D45DFB938C3EFACD1100" ID-GBIF-Occurrence="3396384343" box="[1118,1293,1340,1376]" collectionCode="MOR" pageId="7" pageNumber="240" specimenCode="MOR 980">MOR 980</materialsCitation>
, 
<materialsCitation id="3B63113DE375D45DFAE98C3EFA0B1100" ID-GBIF-Occurrence="3396384328" box="[1316,1483,1340,1376]" collectionCode="BHI" pageId="7" pageNumber="240" specimenCode="BHI 4812">BHI 4812</materialsCitation>
, and 
<materialsCitation id="3B63113DE375D45DF9E38C3EF9101100" ID-GBIF-Occurrence="3396384323" box="[1582,1744,1340,1376]" collectionCode="BHI" pageId="7" pageNumber="240" specimenCode="BHI 6230">BHI 6230</materialsCitation>
all sport extra perforations, thought to be evidence of face biting (
<bibRefCitation id="EF9A6691E375D45DF98D8C73FD2811A9" author="Tanke, D. H. &amp; Currie, P. J." editor="Perez-Moreno, B. P &amp; Holtz, T. J. &amp; Sanz, J. L. &amp; Moratalla, J." journalOrPublisher="Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon" pageId="7" pageNumber="240" pagination="167 - 184" part="15" refId="ref6301" refString="Tanke, D. H., and Currie, P. J. 1998. Elead-biting behavior in theropod dinosaurs: paleopathological evidence. P. 167 - 184 in Perez-Moreno, B. P, Holtz, T. J., Sanz, J. L., and Moratalla, J. (eds.). Aspects ofTheropod Paleobiology. Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon, 15." title="Elead-biting behavior in theropod dinosaurs: paleopathological evidence" type="book" volumeTitle="Aspects ofTheropod Paleobiology" year="1998">Tanke and Currie 1998</bibRefCitation>
; Larson 2001; Rothschild and Molnar this volume).
</paragraph>
<paragraph id="8BB41B60E375D45DFD2C8CE2FCD41312" blockId="7.[658,1904,165,2708]" pageId="7" pageNumber="240">Dentaries: The dentaries each have 13 alveoli. They also both exhibit pathologies. The right dentary has a healed puncture, and a healed puncture and tear near the posterior margin. There is also a 4.5 cm by 1.5 cm erosion located 5 cm below the fifth alveolus. This erosion on the right dentary shows no healing or spongy bone and may be evidence of a fungal infection. The left dentary shows a 4 cm by 1 cm abrasion located 5 cm below the 13th alveolus. This abrasion does show7 evidence of new bone growth.</paragraph>
<paragraph id="8BB41B60E375D45DFD2D8E87FC0E1E48" blockId="7.[658,1904,165,2708]" pageId="7" pageNumber="240">
The skull of 
<taxonomicName id="4C0B60E3E375D45DFC048E87FB7D13C9" authorityName="Osborn" authorityYear="1905" box="[969,1213,1925,1961]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="240" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
(and other theropods) are often reconstructed with the symphysis of the lower jaws widely separated, sometimes by as much as 30 cm (i.e., 
<materialsCitation id="3B63113DE375D45DFB988EF2FA851C74" ID-GBIF-Occurrence="3396384339" box="[1109,1349,2032,2068]" collectionCode="LACM" pageId="7" pageNumber="240" specimenCode="LACM 23844">LACM 23844</materialsCitation>
). This seems to be an attempt to force the teeth of the lower jaws to occlude with those of the upper. Of course, theropods are not crocodiles, and their teeth did not occlude in life. Rather, the teeth of the lower jaws pass medial to those of the upper jaws as the jaws close. Evidence for this is presented by 
<bibRefCitation id="EF9A6691E375D45DFA3581C7F91D1C89" author="Molnar, R. E." box="[1528,1757,2245,2281]" journalOrPublisher="Palaeontographica Abteilung A" pageId="7" pageNumber="240" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991</bibRefCitation>
, p. 143), who noted, “The medial face of the maxilla bears a number of shallow depressions... assumed to have accommodated the tooth crowns [of the dentary] when the mouth was closed, as in alligatoroids.” Thus, the symphysis of the lower jaws were joined by a ligamentous attachment that allowed only limited movement for the purpose of absorbing the shock of teeth hitting bone.
</paragraph>
<paragraph id="8BB41B60E375D452FD2E833CFE5F15D3" blockId="7.[658,1904,165,2708]" lastBlockId="8.[175,1418,181,2084]" lastPageId="8" lastPageNumber="241" pageId="7" pageNumber="240">
Splenials: Both splenials of 
<materialsCitation id="3B63113DE375D45DFB14833BFAB71E3D" ID-GBIF-Occurrence="3396384304" box="[1241,1399,2617,2653]" collectionCode="BHI" pageId="7" pageNumber="240" specimenCode="BHI 3033">BHI 3033</materialsCitation>
are preserved as separate elements showing the lateral surface which articulates against the dentary. 
<emphasis id="B97FC772E375D45DF89E836CF8AD1EF2" box="[1875,1901,2670,2706]" italics="true" pageId="7" pageNumber="240">A</emphasis>
ridge on the dorsal anterior surface of the splenial fits into and becomes part of the Meckelian groove of Osborn (1912). This seems to indicate that there is no movement between the dentary and splenial. The dorsalmost portion of the splenial covers the coronoid, just posterior to the last alveolus of the dentary.
</paragraph>
<paragraph id="8BB41B60E37AD452FF3688C8FAFB179D" blockId="8.[175,1418,181,2084]" pageId="8" pageNumber="241">
CORONOIDS: Only the right coronoid was recovered, but it was preserved in its entirety. 
<bibRefCitation id="EF9A6691E37AD452FDD988F9FD37167F" author="Molnar, R. E." box="[532,759,507,543]" journalOrPublisher="Palaeontographica Abteilung A" pageId="8" pageNumber="241" pagination="137 - 176" part="217" refId="ref6056" refString="Molnar, R. E. 1991. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176" title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1991">Molnar (1991</bibRefCitation>
, p. 155) accepted Osborn’s (1912) interpretation of the coronoid as “a small triangular plate laying at the antidorsal angle of the Meckelian fossa.” However, because the coronoid passes behind the splenial in 
<materialsCitation id="3B63113DE37AD452FDE18B9BFCC916DD" ID-GBIF-Occurrence="3396384374" box="[556,777,665,701]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761" pageId="8" pageNumber="241" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
(and in other articulated 
<taxonomicName id="4C0B60E3E37AD452FB768B9BFF231693" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FB768B9BFF231693" italics="true" pageId="8" pageNumber="241">Tyrannosaurus</emphasis>
</taxonomicName>
lower jaws), Osborn’s interpretation was incorrect. The coronoid actually continues behind the splenial and overlaps the interdental plates of the dentary in what Osborn (1912) called the supradentary. 
<bibRefCitation id="EF9A6691E37AD452FB808A3BFAF2173D" author="Currie, P. J." box="[1101,1330,825,861]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="8" pageNumber="241" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie (2003)</bibRefCitation>
, noting that the splenial and supradentary were joined, and called the entire structure a “coronoid-superdentary” in his description of 
<emphasis id="B97FC772E37AD452FBB68AA1FA4117A7" box="[1147,1409,931,967]" italics="true" pageId="8" pageNumber="241">
<taxonomicName id="4C0B60E3E37AD452FBB68AA1FABC17A7" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[1147,1404,931,967]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
.
</emphasis>
<bibRefCitation id="EF9A6691E37AD452FF7C8ADBFDAA179D" author="Hurum, J. H. &amp; Sabath, K." box="[177,618,985,1021]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="8" pageNumber="241" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath (2003)</bibRefCitation>
referred to it as a “superdentary/coronoid.”
</paragraph>
<paragraph id="8BB41B60E37AD452FF368D0DFCD11C44" blockId="8.[175,1418,181,2084]" pageId="8" pageNumber="241">
<bibRefCitation id="EF9A6691E37AD452FF368D0DFE241053" author="Brochu, C. A." box="[251,484,1039,1075]" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="8" pageNumber="241" pagination="1 - 138" part="7" refId="ref5457" refString="Brochu, C. A. 2003. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology Memoir 7: 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003)</bibRefCitation>
provides a long discussion about whether this element is a single bone (the coronoid) or 2 separate bones (the coronoid and supradentary) that fused during ontogeny. His argument that no immature tyrannosaur individuals display these as separate elements seems to indicate that this is a single element and there is no supradentary. Certainly in 
<taxonomicName id="4C0B60E3E37AD452FB5A8DE1FA431167" authorityName="Osborn" authorityYear="1905" box="[1175,1411,1251,1287]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FB5A8DE1FA431167" box="[1175,1411,1251,1287]" italics="true" pageId="8" pageNumber="241">Tyrannosaurus</emphasis>
</taxonomicName>
(
<materialsCitation id="3B63113DE37AD452FF718C1BFEA1115D" ID-GBIF-Occurrence="3396384324" box="[188,353,1305,1341]" collectionCode="BHI" pageId="8" pageNumber="241" specimenCode="BHI 3033">BHI 3033</materialsCitation>
, 
<materialsCitation id="3B63113DE37AD452FEB58C1BFDDD115D" ID-GBIF-Occurrence="3396384357" box="[376,541,1305,1341]" collectionCode="BHI" pageId="8" pageNumber="241" specimenCode="BHI 4812">BHI 4812</materialsCitation>
, 
<materialsCitation id="3B63113DE37AD452FDF98C1BFD30115D" ID-GBIF-Occurrence="3396384332" box="[564,752,1305,1341]" collectionCode="MOR" pageId="8" pageNumber="241" specimenCode="MOR 1125">MOR 1125</materialsCitation>
, 
<materialsCitation id="3B63113DE37AD452FCCA8C1BFBD8115D" ID-GBIF-Occurrence="3396384306" box="[775,1048,1305,1341]" collectionCode="FMNH" pageId="8" pageNumber="241" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
, etc.), 
<taxonomicName id="4C0B60E3E37AD452FB468C1BFE5E1112" authority="(Currie 2003)" baseAuthorityName="Currie" baseAuthorityYear="2003" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FB468C1BFA43115D" box="[1163,1411,1305,1341]" italics="true" pageId="8" pageNumber="241">Daspletosaurus</emphasis>
(
<bibRefCitation id="EF9A6691E37AD452FF718C4CFE531112" author="Currie, P. J." box="[188,403,1358,1394]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="8" pageNumber="241" pagination="191 - 226" part="48" refId="ref5624" refString="Currie, P. J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologia Polonica 48 (2): 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003</bibRefCitation>
)
</taxonomicName>
, 
<taxonomicName id="4C0B60E3E37AD452FE748C4CFD491112" authorityName="Cope" authorityYear="1866" box="[441,649,1358,1394]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
(TCM 2001.89.1), and 
<taxonomicName id="4C0B60E3E37AD452FBFB8C4CFAC01112" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" box="[1078,1280,1358,1394]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FBFB8C4CFAC01112" box="[1078,1280,1358,1394]" italics="true" pageId="8" pageNumber="241">Tarbosaurus</emphasis>
</taxonomicName>
(
<emphasis id="B97FC772E37AD452FAD28C4CFA451112" box="[1311,1413,1358,1394]" italics="true" pageId="8" pageNumber="241">ZPAL</emphasis>
MgDI/4; 
<bibRefCitation id="EF9A6691E37AD452FE828C80FD2C11C6" author="Hurum, J. H. &amp; Sabath, K." box="[335,748,1410,1446]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="8" pageNumber="241" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath 2003</bibRefCitation>
) the “coronoid-supradentary” is a single bone. The same is true for other theropods like 
<taxonomicName id="4C0B60E3E37AD452FC018CBAFB3111BC" authorityName="Stovall &amp; Langston" authorityYear="1950" box="[972,1265,1464,1500]" class="Reptilia" family="Carcharodontosauridae" genus="Acrocanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FC018CBAFB3111BC" box="[972,1265,1464,1500]" italics="true" pageId="8" pageNumber="241">Acrocanthosaurus</emphasis>
</taxonomicName>
(NCSM 14345) and the allosauroid Sinraptor (
<bibRefCitation id="EF9A6691E37AD452FCDB8CEFFB451271" author="Currie, P. J. &amp; Zhao, X. J." box="[790,1157,1517,1553]" journalOrPublisher="Canadian Journal of Earth Sciences" pageId="8" pageNumber="241" pagination="2007 - 2081" part="30" refId="ref5658" refString="Currie, P. J., and Zhao, X. J. 1993. A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjian, People's Republic of China. Canadian Journal of Earth Sciences 30: 2007 - 2081" title="A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjian, People's Republic of China" type="journal article" year="1993">Currie and Zhao 1993</bibRefCitation>
, p. 2055) where this element is illustrated as a “ceratobranchial.” Even in the prosauropod 
<taxonomicName id="4C0B60E3E37AD452FF7D8F55FEBC121B" authorityName="Meyer" authorityYear="1837" box="[176,380,1623,1659]" class="Reptilia" family="Plateosauridae" genus="Plateosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FF7D8F55FEBC121B" box="[176,380,1623,1659]" italics="true" pageId="8" pageNumber="241">Plateosaurus</emphasis>
</taxonomicName>
it has been interpreted by 
<bibRefCitation id="EF9A6691E37AD452FCEB8F55FBC6121B" author="Galton, P. M." box="[806,1030,1623,1659]" editor="Weishampel, D. &amp; Dodson, P &amp; Osmolska, H." journalOrPublisher="University of California Press, Berkeley" pageId="8" pageNumber="241" pagination="320 - 344" refId="ref5745" refString="Galton, P. M. 1990. Basal Sauropodomorpha. P. 320 - 344 in Weishampel, D., Dodson, P, and Osmolska, H. (eds.). The Dinosauria. University of California Press, Berkeley." title="Basal Sauropodomorpha" type="book chapter" volumeTitle="The Dinosauria" year="1990">Galton (1990)</bibRefCitation>
as a single element: the coronoid. Likewise, Romer illustrates the medial aspect of the lower jaws of 
<taxonomicName id="4C0B60E3E37AD452FF7D8FC3FDD11285" authorityName="Cope" authorityYear="1896" box="[176,529,1729,1765]" class="Reptilia" family="Captorhinidae" genus="Labidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FF7D8FC3FE4C1285" box="[176,396,1729,1765]" italics="true" pageId="8" pageNumber="241">Labidosaurus</emphasis>
(1956a)
</taxonomicName>
, 
<emphasis id="B97FC772E37AD452FDE88FC3FB941285" box="[549,1108,1729,1765]" italics="true" pageId="8" pageNumber="241">
<taxonomicName id="4C0B60E3E37AD452FDE88FC3FD7A1285" authority=", Diadectes, Bradysaurus, Python" authorityName="Amalitzki" authorityYear="1921" box="[549,698,1729,1765]" class="Amphibia" family="Kotlassiidae" genus="Kotlassia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Seymouriamorpha" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Kotlassia</taxonomicName>
, 
<taxonomicName id="4C0B60E3E37AD452FD028FC3FCB11285" authority=", Diadectes, Bradysaurus, Python" authorityName="Amalitzki" authorityYear="1921" box="[719,881,1729,1765]" class="Amphibia" family="Kotlassiidae" genus="Kotlassia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Seymouriamorpha" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Diadectes</taxonomicName>
, 
<taxonomicName id="4C0B60E3E37AD452FC488FC3FB8E1285" authority=", Diadectes, Bradysaurus, Python" authorityName="Amalitzki" authorityYear="1921" box="[901,1102,1729,1765]" class="Amphibia" family="Kotlassiidae" genus="Kotlassia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Seymouriamorpha" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Bradysaurus</taxonomicName>
,
</emphasis>
<taxonomicName id="4C0B60E3E37AD452FBA98FC3FB181285" authority=", Diadectes, Bradysaurus, Python" authorityName="Amalitzki" authorityYear="1921" box="[1124,1240,1729,1765]" class="Amphibia" family="Kotlassiidae" genus="Kotlassia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Seymouriamorpha" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Python</taxonomicName>
, 
<emphasis id="B97FC772E37AD452FB208FC3FF21137B" italics="true" pageId="8" pageNumber="241">
<taxonomicName id="4C0B60E3E37AD452FB208FC3FF1B137B" authority=", Dimetrodon" authorityName="Lydekker" authorityYear="1889" class="Reptilia" family="Pliosauridae" genus="Peloneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Plesiosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Peloneustes</taxonomicName>
,
</emphasis>
<taxonomicName id="4C0B60E3E37AD452FF218FF5FE72137B" authority=", Dimetrodon" authorityName="Lydekker" authorityYear="1889" box="[236,434,1783,1819]" class="Reptilia" family="Pliosauridae" genus="Peloneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Plesiosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Dimetrodon</taxonomicName>
, and 
<taxonomicName id="4C0B60E3E37AD452FDC48FF5FCB3137B" authorityName="Cope" authorityYear="1882" box="[521,883,1783,1819]" class="Reptilia" family="Edaphosauridae" genus="Edaphosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pelycosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">
<emphasis id="B97FC772E37AD452FDC48FF5FD33137B" box="[521,755,1783,1819]" italics="true" pageId="8" pageNumber="241">Edaphosaurus</emphasis>
(1956b)
</taxonomicName>
as all possessing a coronoid with a thinning anterior projection that passes over the medial aspect of much of the tooth row—the same situation seen in theropods. It is for these reasons that the “coronoid-supradentary” should simply be referred to as the coronoid. The coronoid for 
<materialsCitation id="3B63113DE37AD452FDF98EC9FD0F138F" ID-GBIF-Occurrence="3396384318" box="[564,719,1995,2031]" collectionCode="BHI" pageId="8" pageNumber="241" specimenCode="BHI 3033">BHI 3033</materialsCitation>
closely resembles that of 
<emphasis id="B97FC772E37AD452FBBA8EC9FA84138F" box="[1143,1348,1995,2031]" italics="true" pageId="8" pageNumber="241">
<taxonomicName id="4C0B60E3E37AD452FBBA8EC9FAFE138F" authorityName="Maleev" authorityYear="1955" baseAuthorityName="Hurum and Sabath" baseAuthorityYear="2003" box="[1143,1342,1995,2031]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
,
</emphasis>
described by 
<bibRefCitation id="EF9A6691E37AD452FE978102FCCD1C44" author="Hurum, J. H. &amp; Sabath, K." box="[346,781,2048,2084]" journalOrPublisher="Acta Palaeontologia Polonica" pageId="8" pageNumber="241" pagination="161 - 190" part="48" refId="ref5818" refString="Hurum, J. H., and Sabath, K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologia Polonica 48: 2, 161 - 190" title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath (2003)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C31148EBE37AD453FA1F819CF984156C" lastPageId="9" lastPageNumber="242" pageId="8" pageNumber="241" type="discussion">
<paragraph id="8BB41B60E37AD452FA1F819CF96A1CB0" blockId="8.[1490,1706,2206,2256]" box="[1490,1706,2206,2256]" pageId="8" pageNumber="241">Conclusion</paragraph>
<paragraph id="8BB41B60E37AD453FF6281A5F984156C" blockId="8.[175,1416,2208,2679]" lastBlockId="9.[645,1880,174,268]" lastPageId="9" lastPageNumber="242" pageId="8" pageNumber="241">
The disarticulated skull ofTyrannosaurus rex specimen known as 
<materialsCitation id="3B63113DE37AD452FB2481A5FA411CAB" ID-GBIF-Occurrence="3396384369" box="[1257,1409,2215,2251]" collectionCode="BHI" pageId="8" pageNumber="241" specimenCode="BHI 3033">BHI 3033</materialsCitation>
preserves many details not seen in other specimens. A close examination reveals details of the air sac system that invade or leave its impressions on many of the bones of the skull. Pathologies—evidence of disease or healed injuries—found on some of the cranial elements provide insight into behavior, such as intraspecific combat (face biting) and the commonality of certain pathogens such as osteomyelitis. And finally, the exquisite preservation of the cranial joint surfaces on 
<materialsCitation id="3B63113DE37AD452FCED831EFC7F1E20" ID-GBIF-Occurrence="3396384355" box="[800,959,2588,2624]" collectionCode="BHI" pageId="8" pageNumber="241" specimenCode="BHI 3033">BHI 3033</materialsCitation>
opens the door to a more complete analysis of the capacity for cranial kinesis in 
<taxonomicName id="4C0B60E3E37AD452FB818352FABC1E14" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1100,1404,2640,2676]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="241" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
Limited discussions of some aspects cranial kinesis in 
<taxonomicName id="4C0B60E3E37BD453F9DD89B1F9A914B7" authorityName="Osborn" authorityYear="1905" box="[1552,1641,179,215]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="242" phylum="Chordata" rank="species" species="rex">
T. 
<emphasis id="B97FC772E37BD453F9F789B1F9A914B7" box="[1594,1641,179,215]" italics="true" pageId="9" pageNumber="242">rex</emphasis>
</taxonomicName>
may be found in 
<bibRefCitation id="EF9A6691E37BD453FD7F89EAFBBC156C" author="Larson P. L. &amp; Donnan, K." box="[690,1148,232,268]" journalOrPublisher="Invisible Cities Press, Montpelier, VT" pageId="9" pageNumber="242" refId="ref6014" refString="Larson P. L., and Donnan, K. 2002. Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life. Invisible Cities Press, Montpelier, VT" title="Rex Appeal: The Amazing Story of Sue, the Dinosaur that Changed Science, the Law and Aly Life" type="book" year="2002">Larson and Donnan (2002)</bibRefCitation>
and Larsson (this volume).
</paragraph>
</subSubSection>
</treatment>
</document>